chromosome studies in lamiaceae -...
TRANSCRIPT
CHROMOSOME STUDIES IN LAMIACEAE
The dicotyledonous family Lamiaceae is a very large natural group of aromatic
and medicinal plants (Morton, 1962). Lindley (1830) was the first to propose
the name, *Lamiaceae' and later on in 1935 the International Code of Botanical
Nomenclature approved that it can be used as an alternative to Labiatae. The
name .Labiatael, coined by de Jussieu (1789) originates from the Latin noun
labium (lip), referring to the bilipped corolla of their zygomorphic flowers,
a salient feature of the family.
Lamiaceae enfolds 220 genera (Hedge, 1992) and almost 5000 species (Mabberley,
1987), with a very wide range of morphological diversity. The historical
background of Lamiaceae is of much importance with regard to it's phylogeny.
The ancestors of the family might have been originated in the tertiary
tropical forests (Chengyih and Hsiwen, 1982). The evolution and specialization
of the family began in the ancient Mediterranean region. The phylogeny of the
family is detailed just as the centres of origin. Zeven and Zhukovsky (1975)
proposes nine different gene centres, which formed cradles of Lamiaceae
cultivation as, the Chinese-Japanese centre, the Indo-Chinese-Indonesian
centre, the Australian centre, the Hindustani centre, the Central Asian
centre, the Near Eastern centre, the Mediterranean centre, the European
Siberian centre and the African centre.
Majority of these plants are annual and perennial herbs, cosmopolitan, growing
under a great variety of soil and climate and constitute a scrub type of
vegetation (Simpson and Conner - Ogorzaly, 1986). Though the members have a
global distribution (Epling, 1940b; Harley and Reynolds, 1992), the prime
centre of concentration lies in the Mediterranean basin and Asia Minor, both
in regards the number of genera and species (Rivera Nunez and Obon de Castro,
1992). The other main centres of diversity include Europe, Middle East,
Tropical and Central Africa, South Africa, Asia, Australasia, North America,
Arctic and Antartic regions (Morley, 1970).
Members of Ocimeae flourishes in tropics and subtropics, represented in both
hemispheres (Rendle, 1925) by 32 genera and 970 species (Morton, 1962).
Mentheae includes the largest group of plants among Lamiaceae, which are
cosmopolitan in distribution and posses 42 genera and about 540 species (EI-
Gazzar and Watson, 1967). The various taxa of Salveae are distributed in the
temperate and warmer regions of both hemispheres with 10 genera and more than
738 species (Bailey, 1960). The aromatic herbs of Nepeteae are concentrated in
subtropical and temperate Asia (Gill, 1979) and are represented by 9 genera
and 393 species (Willis, 1973). Lanieae consists of a large group of plants,
cosmopolitan in distribution with about 37 genera and about 850 species
(Craner, 1981). Members of Prasieae have a limited distribution, chiefly in
tropical Asia and the Ha~iai Islands (Rendle, 1925), represented by 5 genera.
(Bentham 1876). Prostanthereae includes a small group of plants confined to
Australia and Tasmania (Lawrence, 1951), represented by a few genera and 200
species (Conn, 1992). The various plants of Ajugeae are cosmopolitan in
distribution (Lawrence, 1951) with about 8 genera (Cantino, et al. 1992). In
India, the family is represented by about 69 genera and 425 species of which
261 are endemic. The temperate Himalayas, the Deccan, North Western India and
South India are the chief centres of concentration. South India is one of the
major centres of distribution since it support over 139 species (Mukherjee,
1940).
The Lamiaceae members are characterized by their peculiar morphology and
aromaticity that they have been well recognized for centuries and is one of
the oldest families known (Swingle, 1946).
The family Lamiaceae is of great importance from a commercial standpoint
(Heywood, 1978). Besides numerous aroma chemicals, perfume products,
medicinals, pot herbs and honey pasturage, the family yield a number of
choice ornamentals (Stanford, 1937). Many members of Lamiaceae find use in
cosmetic industry (Fleisher and Fleisher, 1991 b; Knaut and Richtler, 1990),
for the preparation of detergents, fabric conditioning mixtures (Behan and
Clements, 1990; Ito and Kikuchi, 1991; Nishida, 1991), deodorants and bath
preparations [Kawasaki and Tsunakawa, 1990; Nishida, 1990; Ueyama, et al.,
1990; Morgan and Blagdon, 1991) long lasting fragrant preparations and
perfumes (Tsukasa, 1990; Yoshihara, et al., 1990; Ikimoto and Kuwaori, 1991;
Okada, et al., 1991), herbal preparations (Muller, 1990, 199l),non-irritating
cleansers (Morganti, 1990), hairgels (Lochhead, 1990) and in baking industry
(Bassiouny, et al., 1990; Economou, et al., 1991). Some species find use in
the preparation of dentifrices, mouthwashes, toothpowders, chewing gum and
lozenges, which act as anti-plaque and anti-tartar agents of teeth (Anonymous,
1991; Reuss, 1991; Mori and Yokosuka, 1991). Antifungal and antibacterial
activities have been reported on many species. (Gangrade, et al., 1989a,
1989b) This property is due to their volatility and ephemeral nature (French,
1985) and is of much importance since herbal fungicides are non-toxic (Fawcett
and Spencer, 1970) and easily bio-degradable (Mahadevan, 1982).
The position affinity and supra-familial relationship of the family Lamiaceae
in different systems of classification has been a subject of considerable
debate from time immemorial. Linnaeus (1737) placed the family under 2
classes. Didynamia and Tetra dynamia. However de Jussieu (1789). Gray (1858)
and Eichler (1875-1878) grouped them together under Sympetalae. Bentham and
Hooker (1862-1883) classified Lamiaceae in the order Lamiales, followed by
Bessey (1915). Hutchinson (19261, Cronquist (19681, Takhtajan (1969) and
Dahlgren (1975). On the basis of morphological parameters, Engler and Prantle
(1888-1897) placed Lamiaceae in the order Tubiflorae. Later on Hallier (1905).
Wettstein (19241, Rendle (1925) and Pulle (1938) followed the same pursuit.
June11 (1934) and Erdtman (1945) established their systems of classification
on the basis of evidence from gynoecial and pollen characters. In a more
recent system of classification proposed by Goebel (1984) Lamiaceae is placed
in the order Labiatiflorae. Recent research suggests that the family Lamiaceae
is polyphyletic (Abu-Asab and Cantino, 1989, 1992; Cantino, 1992 a, b).
Bentham (1832-1836, 1848, 1876), and Briquet (1895-1897) have made major
contributions towards the infra-familial classification of the family. Later
on modified systems have been proposed by Hillson (1959), Melchior (1964).
Wunderlich (1967), El-Gazzar and Watson (1970a), Dahlgren (1980). Taktajan
(1980), Thorne (1983), Cantino and Sanders (1986) as well as Goldberg (1986).
Bentham (1876) Sub divided Lamiaceae into eight tribes, which are Ocimoideae
(*Ocimeae), Satureineae (*Mentheae), Monardeae (*Salvieae), Nepeteae, Lamieae
(*Stachydeae) Prasieae, Prostanthereae and Ajugoideae (*Ajugeae).
The familial relationship of Lamiales have been evaluated by Cantino (19821,
Cantino et al. (1992) Cronquist (1981) and Thorne (1976) and supra-ordinal
relationships by Lu (1990). The family Lamiaceae is readily distinguished from
all the other families except Verbenaceae by the distinctive gynoecium
(Lawrence, 1951). It is considered to be closely resembling Verbenaceae in
having similar vegetative as well as floral characteristics. The presence of
* Correct names proposed by International Code of Botanical Nomenclature
(Stafleu, et al., 1978; Greuter, 1981).
these intergrading characters make it difficult, if not impossible to separate
all members of one family from the other by any single charac.ter or a
combination of characters (Rendle, 1925). Thus the family Verbenaceae is
phylogenetically related to Lamiaceae (Bessey, 1897). However, Hutchinson
(1924) considered that the two are unrelated, but the two families seem rather
similar because both of them represent climaxes of evolution in their
respective groups. The complicated floral structure among the gamopetalous
families, corroborating with various adaptations for entomophily show that
Lamiaceae can be considered undoubtedly as the culmination of the sympetalous
herbaceous line of Herbaceae (Hutchinson, 1926).
In addition to this Lamiaceae resembles Scrophulariaceae and Acanthaceae by
showing more or less parallel floral structure (Hutchinson, 1924), as well as
the occasional quadrangular nature of stem and opposite phyllotaxy (Rendle,
1925). However according to Evans (1936) Lamiaceae differs from both by the
presence of verticillaster inflorescence, gyno-basic style and the 4-lobed
nature of the ovary as well as fruit.
Lamiaceae also bears relationship with the Boraginaceae, but is set apart by
the technical character of the ovule. When the micropyle is inferior and raphe
inward in Lamiaceae, it is just the reverse in Boraginaceae (Lawrence, 1951).
More over Lamiaceae possess fewer number of stamens than the corolla lobes
(Evans, 1936). Thus Lamiaceae is entitled to a rather high rank among the
Metachlamydeae by reason of their irregularity, tetracyclic condition, reduced
number of stamens as well as carpels and highly specialized type of fruit
(Haupt, 1946).
The perennial growth habit, simple leaves, heterogenous medullary rays,
bisexual hypogynous flowers, anatropous ovules and entomophilous pollination
are some of those primitive features shown by the family (Coulter and
Chamberlain, 1903, Eames, 1961; Metcalfe and Chalk, 1950; Takhtajan, 1969).
The herbaceous nature, exstipulate, opposite or whorled leaves, small
zygomorphic flowers, bilabiate calyx and corolla showing cohesion, reduced
number of stamens, exhibiting epipetaly, nearly monothecous condition of
anthers, bicarpellary syncarpous pistil showing axile placentation, reduction
of the number of ovules to one in each loculus, simple fruits, non-endospermic
seeds and presence of small sized vessels and fibres with simple pits are some
of the advanced characters exhibited by the family (Bessey, 1893; Cronquist,
1968; Erdtman, 1952; Hallier, 1905; Metcalfe and Chalk, 1950; Nageli, 1884;
Takhtajan, 1959; Bailey and Tupper, 1918).
Some of the common genera found in India include Ocimum L.; Acrocephalus
Benth.; Orthosiphon Benth.; Plectranthus L'Herit.; Coleus Lour.; Anisochilus
WalL, Hyptis Jacq.; Pogostenon Desf.; Eusteralis Rafin.; Mentha L.; Salvia L.;
Anisomeles R.Br.; Leucas R.Br.; Leonotis R.Br. and Teucrium L.
Ocimum L., the common *Basil' derives its name from Greek, where 'ozo' means
smell (Hereman, 1868). It possess 160 species with the maximum members
originating from the tropical forests of Africa (Sobti and Pushpangadan,
1982). Other main centres of diversity include South America and Asia
(Krishnamoorthy, (1989). The medicinal and pharmacological properties of this
genus are very well described, in the treatises *Pharmacographia Indica'
(Dymock, et al; 1890) and *Indian Materia Medica' (Nadkarani, 1976 a, 1976 b).
It is well known in perfumery, flavour and cosmetic industries (Kirtikar and
Basu, 1935; Guenther, 1952; Dastur 1962).
The plant of little beauty, Acrocephalus Benth, is named after .akros1 -
summit and *kephalel - head; on account of the flowers being on top of the
branches (Hereman, 1868). This tropical genus, with about 100 species
(Santapau and Henry, 1973) is distributed in the warmer parts of Asia and
Africa (Hooker, 1885). Some species are used in folk medicines (Chopra, et
al., 1956).
The generic name Or-thosiphon Benth., originates from 'orthos' - straight and
-siphon1 - tube; refers to the straight corolla tube (Bailey, 1960). It is
represented by more than 100 species (Santapau and Henry, 1973), mainly annual
or perennial herbs and are distributed in the tropics from Africa to Australia
(Bailey, 1960). Some species are employed in Ayurvedic and Unani medicines
(Uphof, 1959).
Plectranthus L' Herit., an Old World genus (Rivera Nunez & Obon de Castro,
1992), is named from *plektronl-Cock's spur and <anthos1-flower; alluding to
the swollen corolla base and hence the common name, .Cock's spur' (Hereman,
1868). A genus with about 300 species (Richardson, 1992) of fleshy herbs and
under shrubs (Bower, 1956) are distributed chiefly in the tropical and
subtropical regions of Africa, and Asia, spreading eastwards to Australia,
Malaysia, Japan and the Pacific region (Bailey, 1960).
The name Coleus Lour., araises from *coleos'-sheath; referring to the
monadelphous stamens (Plowden, 1968). This Old World genus, commonly known as
the -Flame-nettle' (Fyson, 1986) possess 150 species (Morley, 1970), that are
mainly distributed in the Eastern Hemisphere, being especially abundant in
Africa, India, the Malayan Archipelagoes, extending to Australia and the
Pacific islands (~aile~, 1960). Coleus Lour. yields a variety of pot and
garden ornamentals, and plants with edible tubers (Swingle, 1946). Some
members are found to be of very high medicinal value (CSIR, 1986).
Anisochilus Wall., derived its name from *anises' unequal and *cheilos' - lip;
with regard to the zygomorphic corolla of the flowers (Hereman, 1868). It is a
cosmopolitan genus and holds about 20 species Cramer, 19811, distributed in
India and Tropical Africa (Fyson, 1986). Medicinal and pharmacological
properties are enumerated in -Pharmacographia Indica' (Dymock et al., 1890)
Hyptis Jacq. is named after .hyptios' - resupinate; since the limb of the
corolla is turned on its back (Hereman 1868). It is a large genus with as many
as 400 species and occur in different parts of the world (Richardson, 1992),
the chief centre of concentration being America (Hooker, 1885). The medicinal
properties are revealed by Chopra, et al. (1956).
The name Pogostemon Desf., is originated from -pogon'-beard and .stemon' -
stamen; owing to the tuft of hairs present in the stamens (Bailey, 1960). The
genus comprises a group of aromatic herbs and shrubs (Bower, 1956) with about
40 species (Cramer, 1981) and is mainly distributed in India and Eastern Asia
(Fyson, 1986). Many species are described in *Pharmacographia Indica' for
their medicinal qualities (Dymock, et al., 1890).
Eusteralis Rafin. is a very close ally of Pogostemon Desf. It is a woddy
aromatic herb very common on the rocky slopes, growing in the crevices. 21
species have been reported so far in this genus (Cramer, 1981).
The name Mentha L., bestowed upon by the Greek philosopher Theopharastus
(Jeans, 1973) after *Menthael, Latin name of a nymph; due to the aquatic
habitat it prefers (coats, 1968). It includes several perennial herbaceous
plants (Ranade, 1982), thriving very well in marshy places and propagate by
means of runners or stolons (Stanford, 1937). It is commonly called as mint
(Gray, 1858). Index Kewensis (1893 - 1947) lists about 900 binomials widely
distributed in the temperate and subtropical regions of the World. However,
Harley and Brighton (1977) estimated that there are probably some 25 species.
The medicinal and aromatic properties of many species are described in
'Materia Indica' (Ainslie, 1826) and 'Pharmacographia Indica' (Dymock, 1890).
The name -a L., is derived from the Latin word 'salvo' which means-I save;
referring to the medicinal qualities (Plowden, 1968). The 'common sage' (Gray,
1858) is the most primitive genus of the family although L. is
considered by many authors (Fujita, 1970). This genus is the largest in the
family Lamiaceae (Morley, 1970), with about 1000 species (Estilai, et al,
1990) and widely distributed in the temperate and warmer regions. 'Materia
Indica' (Ainslie 1826') gives a detailed account of its medicinal uses.
The genus' Anisomeles R.Br. is named from 'anisos' unequal and 'melos' -
member; in collaboration with the corolla character (Hereman, 1868). This
small genus with 7 species (Cramer, 1981) is distributed in tropical and
subtropical Asia and Australia (Hooker, 1885). Medicinal properties are
described by Kirtikar and Basu (19351, Uphof (1959), Dastur, (1962) and CSIR
(1986).
Leucas R.Br. is named after 'leukos' - white; in accordance with the downy
white flowers (Hereman, 1868). The genus commonly known as 'Dead-nettle'
(Fyson, 1986) includes wooly, villous pubescent herbs or shrubs with about 60
species (Cramer, 1981) and is concentrated in Asia and Africa (Fyson, 1986).
Medicinal and pharmacological notes are given by Chopra, et al. (1956).
The name Leonotis R.Br. is derived from 'leon' - lion and 'ous' - ear'
referring to the resemblance of the flower to lion's ear. ( Plowden, 1968:. It
consists of annual or perennial herbs or shrubs with about 20 species found
mainly in tropical and South Africa (~ailey, 1960). Medicinal aspects are
mentioned in 'Pharmacographia Indica' (Dymock, et al., 1890).
Teucrium L., is named after -Teucer' of Greek mythology, who discovered it's
medicinal uses (Plowden, 1968). The genus is commonly known as *Germander1
(Gray, 1858). Many of its species are natives of the Mediterranean (Morley,
1970) and chiefly distributed in Temperate and Southern Europe (Fyson, 1986).
The genus consists of more than 100 species (Gleason, 1963).
During the past few decades, the cytological data of many families have been
widely used in comparative and evolutionary studies of various taxa. In
plants, the cytological data of taxonomic importance is mainly pertaining to
the chromosome number and structure during mitosis. Chromosome number will
serve as useful taxonomic markers to distinguish between different taxa.
Additionally it can also solve the problems of dispute between different taxa.
Therefore, these factors are used as classificatory criteria in the same
manner as the morphological characters since the chromosomes have a direct
relation to the genetic system of which they are an integral part (Den Hartog,
et al. 1979).
Sufficient information is available on the Indian members of Lamiaceae by
virtue of the major cytological surveys conducted by Gill (1970, 1971a, 1971b,
1971~) Mehra and Gill (1972), Bhattacharya (1976, 1978a, 1978b) Vij and
Kashyap (1976b) Singh and Sharma (1981a, b, 1982) Vembu and Sampathkumar
(1981), Bir and Saggoo (1984, 1985), Saggoo and Bir (1983a, 1985, 19861, Kundu
and Sharma (1988a). However, almost all these cytological studies have
concentrated mainly on the determination of chromosome number and provided
scarcely any data on chromosome morphology and structure in detail.
Karyomorphological studies are of paramount importance as they often provide
authentic information pertaining to chromosome structure, number and in
general their gross morphology (Darlington and La Cour, 1970). Detailed
studies pertaining to chromosome length, disparity index of chromosomes,
variation coefficient among chromosome complement and total forma percentage
are necessary in order to throw light on the phylogenetic relationships among
the taxa of flowering plants, since chromosome number tend to exhibit a
greater consistency than any other character. (Yoshikane and Naohiro, 1991).
Moreover,Jones (1978) pointed out that in order to determine the polarity of
karyotype evolution, it is useful to integrate the study of karyotype
morphology with other aspects of the organism, like chemical constitution.
Therefore in the present investigation karyomorphological analysis is
conducted on fifty four members of South Indian Lamiaceae with the aid of "---- " ,-..
1 .' $8 &. d the improved techniques (Sharma and Sharma, 1980) in order to .kt& . yp
number, structure and behaviour of their
correlation between the chromosome
quantitative changes of their essential oil
Fifty four taxa of Lamiaceae has been collected from different parts of South
India, situated between the north latitudes 8 and 20 and the east longitudes
74 and 86. (vide. Table 1; Fig.1).
Table 1 --
List of aerbers investigated
S1. No.
1.
2.
3.
4.
5 .
6.
7 .
8.
9.
10.
11.
12.
13.
14.
15.
Name of the taxa
Ocimum adscendens Willd.
0- americanum L.(syn 0. canum Sims)
0. basilicum L.var. glabratum Benth.
- 0. basilicum L.var pilosum Benth.
- 0. basilicum L.var. purpurascens Benth.
O, basilicum L.var thyrsiflorum Benth.
0. - L. -
O, gratissimum L.var. suavis H0ok.f.
- 0. tenuiflorum L.f.(syn. - 0. sanctum L.) cv. green
- 0. tenuiflorum L.f.cv. purple
O, tenuiflorum L.f. cv.purple-green
- 0. tenuiflorum L.f.var. hirsuta H0ok.f.
Acrocephalus capitatus Benth.
Orthosiphon glabratus Benth.
& glabratus var. parviflorus Benth.
Locality of collection
Cherthalai
Calicut
Cherthalai
Mysore
Wynad
Kodaikanal
Cochin
Bangalore
Kottayam
Munnar
Cochin
Ooty
Cochin
Kodaikanal
Palani
Altitude in metres (approx) .
Sea Level
45
Sea Level
450
1200
2100
Sea Level
920
80
1050
Sea Level
2290
Sea Level
2100
800
- a. No.
- 16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
37. -
Name of the taxa
0. srandiflorus Bold(syn O, stamineus Benth. ) - cv. lilac
0. grandiflorus Bold cv. white - 0. pallidus Royle - 0. thymiflorus Roth. - Plectranthus nummularius Briq.
P. wightii Benth. - Coleus aromaticus Benth. (syn. amboinicus
Lour. )
C. aromaticus Benth. var. variegata - C. blumei Benth. var. verschaffeltii - C. forskohlii Briq. - C. laciniatus Benth.
C. parviflorus Benth. - C. rehneltianus Berger - C. zeylanicus(Benth.) Cramer - Anisochilus carnosus (L.f.1 Wall.
A. eriocephalus Benth. - Hyptis capitata Jacq.
H. suaveolens (L.) Poit. - Pogostemon benghalensis (Burm.f.)Kuntze
P. heyneanus Benth. - P. purpurascens Dalz. - P. vestitus Benth. -
Palode
loty
Kodaikanal
Ponmudi
Yunnar
Cochin
Palani
Bangalore
Cochin
Trivandrul
Palghat
Idukki
Kottayam
Wynad
Cochin
Munnar
Ponmudi
Palghat
Cochin
ltitude n metres approx) .
180
2290
2100
900
1050
Sea Level
800
920
Sea Level
50
150
1000
80
1200
Sea Level
1050
900
150
Sea Leve:
1 . Ocimum adscendens Willd. (Fig.2a) Annual herb; stem fleshy, erect or
prostrate, quadrangular; leaves somewhat fleshy, broadly serrate, ovate-
obovate, membraneous; inflorescence a lax terminal raceme; flowers small,
S1. No.
38.
39 .
40 .
41 .
42.
43 .
44.
45.
46.
47.
48.
49 .
50.
51.
52.
53 .
54 .
blue; tropical.
Altitude in metres (approx) .
45
2290
1800
Sea Level
450
1050
1000
Sea Level
900
Sea Level
1050
45
1220
800
900
Name of the taxa
Eusteralis quadrifolia (Benth.) Panigrahi
Mentha arvensis L.
- M. rotundifolia (L.) Huds.
M. spicata (L.) Huds.
Salvia coccinea Juss.
& leucantha Cav.
- S. plebeia R.Br.
- S. splendens Ker-Gawl.
Anisomeles indica (L.) Kuntze
- A. malabarica R.Br.
Leucas aspera Spreng.
Z, cephalotes Spreng.
- L. linifolia Spreng.
- L. stricta Benth.
- L. vestita Benth.
Leonotis nepetifolia (L.) Ait.f.
Teucrium plectranthoides Gamble
Locality of collection
Calicut
Ooty
Coonoor
Cochin
Mysore
Munnar
Idukki
Cochin
Ponmudi
Cochin
Munnar
Calicut
Devikolam
Palani
Thekkadi
2. 0. americanum L. (Fig. 3a) 'Hoary basil'; origin - Tropical Africa;
perennial herb; stem 4 - angled, grooved; leaves glabrous, simple,
elliptic, slightly crenate; inflorescence a terminal raceme; flowers
white; tropical.
3. 0. basilicum L.var. glabratum Benth. (Fig. 4a) 'French basil'; origin -
France; perennial herb; stem four angled, woody; leaves ovate, glabrous,
margins serrate, acuminate at apex; inflorescence a long raceme; flowers
yellow; tropical and subtropical.
4. 0. basilicum L. var pilosum Benth. (Fig. 5a) Perennial herb; stem
quadrangular, highly pubescent; leaves ovate, acute and pilose with
glandular hairs, margins sparingly serrate; inflorescence a slender
raceme, axis pilose; flowers white; tropical and subtropical.
5. 0. basilicum L. var. purpurascens Benth. (Fig. 6a) 'Sweet basil'; origin
- Tropical Asia, Africa and Pacific Islands; perennial herb; stems
square, tender parts purple; leaves opposite, petioles long, blades
broad, ovate-acute, coarsely crenate, inflorescence a spike, terminal or
axillary; flowers pink or purple; tropical and subtropical.
6. 0. basilicum L. var. thyrsiflorum Benth. (Fig. 7a) Perennial herb; stem
stout, tetragonous; branchlets glabrascent; leaves elliptic - lanceolate, thick, chartaceous, margin serrate, apex narrowly acute; inflorescence
typical thyrse - like racemes; flowers pink, tropical and subtropical.
7. 0. gratissimum L. (Fig.8a) 'Shrubby basil'; origin-Tropical Africa,
America and India; perennial undershrub; stem 4 - angled, 4 - grooved,
young branches pubescent; leaves large, obovate, acute at apex, cuneate
Fig. 2a
Fig. 5a I!---,, Fig. 6a
- Fig. 4a
I I Fig. 7a
Fig. 10a
at base, crenate-serrate, long-petioled, glabrous except the nerves;
inflorescence a terminal raceme; flowers pale greenish yellow; tropical.
8. 0. gratissinn L. var. suavis H0ok.f. (Fig.9a) Perennial undershrub;
stern-&angular; leaves elliptic-lanceolate, acute, coarsely crenate,
pubescent and gland-dotted on both sides, base cuneate, petioles slender;
inflorescence a raceme; flowers greenish-yellow, pedicels softly hairy;
tropical and subtropical.
9. 0. tenuiflonm L.f. cv. green (Fig. 10a) 'Holy basil'; origin-Tropics of
the Old World; perennial herb; stem woody at base, branches clothed with
soft pubescent hairs, greenish, leaf blade elliptic-oblong, apex obtuse,
margins subterrete, hairy on both surfaces; inflorescence a thyrsus,
slender; flowers in close whorls, purplish-pink; tropical.
10. 0. tenuiflorm L.f. cv. purple (Fig. lla) 'Sacred basil'; perennial herb;
stem woody and purplish; leaves ovate-oblong; crenate-serrate, purple,
hairy; inflorescence a lax raceme of verticils; flowers purple in colour;
tropical.
11. 0. tenuiflonm L.f. cv. purple-green (Fig. 12a) Perennial herb; stem
solid, 4-angled, branched, hairy, woody below and herbaceous above;
leaves simple, opposite and decussate, ovate, serrate, acute, purplish-
green; inflorescence a raceme of thyrses; flowers small, pink; tropical.
12. 0. tenuiflorm L.f. var. hirsuta H0ok.f. (Fig. 13a) Perennial herb; stem
slender, hairy, quadrangular, branched; leaves simple, opposite,
petioled, narrowly ovate, toothed, green; inflorescence a thyrsus, axis
pilose; flowers yellow, small; tropical and subtropical.
13. Acrocephalus capitatus Benth. (Fig. 14a) Annual herb; stem erect,
branching from the base, bluntly 4-angled, glabrous; leaves small,
lanceolate, subacute, coarsely serrate, tapering at the base;
inflorescence a terminal capitate head, subtended by a pair of foliar
bracts; flowers small, pale lilac; tropical.
14. Orthosiphon glabratus Benth. (Fig. 15a) Erect perennial undershrub; stem
glabrous, tetragonous, stout; leaves broadly ovate-cordate, membraneous,
chartaceous, margin deeply crenate, scarcely tomentose on the veins;
inflorescence a lax raceme; flowers medium sized, pinkish white; tropical
and subtropical.
15. 0. glabratus var. parviflorus Benth. (Fig. 16a) Perennial undershrub;
stem glabrous, quadrangular, 4-grooved; leaves broadly ovate, coarsely
serrate, nearly glabrous; inflorescence a short raceme; flowers small,
narrow , pale purplish; tropical.
16. 0. grandiflorus Bold cv. lilac (Fig. 17a) 'Kidney Tea plant' or 'Java
Tea'; origin-Tropical Asia and Malayan Archipelago; erect perennial herb;
stem 4-angled, 4-grooved, stout, purple to green coloured; leaves ovate-
acute, margin coarsely toothed, blade smooth and glabrascent;
inflorescence a large flowered raceme, terminal, whorls crowded; flowers
tubular, pale lilac-blue with long protruding stamens; tropical.
17. 0. grandiflorus Bold cv. uMte(Fig. 18a) Perennial herb; stem
tetragonous, sinused; leaves glabrous, olivegreen; broadly ovate,
crenate-serrate; inflorescence a raceme; flowers large, showy, white with
conspicuous very long filiform stamens; tropical.
Fig. 14a
Fig. 17a
Fig. 12a
Fig-. 18a
Fig . 13a
Fig. 16a
Fig . 19a
18. 0. paJJ.idus Royle (Fig. 19a) A low perennial herb; stem glabrous;
branches spreading; leaves small, ovate, toothed, petiolate, pale green;
inflorescence a small terminal raceme; flowers white; tropical and
subtropical.
19. 0. thymiflorus Roth. (Fig. 20a) Perennial herb; stem 4-angular,
glabrascent; leaves broadly ovate to cordate acute, chartaceous, margin
serrate; inflorescence a terminal raceme; flowers pale purple; tropical.
20. Plectranthus nltmmularius Briq. (Fig.2la) 'Swedish Ivy'; origin-Australia
and Pacific Islands; perennial herb; stem weak, creeping; leaves
leathery, metallic green, waxy, rounded, deeply crenate; inflorescence a
lax spike; flowers white; tropical and subtropical.
21. P. wightii Benth. (Fig.22a) Tall robust; stem scaberulous; leaves broadly
ovate-cordate, crenate-toothed, rigid; inflorescence a large pyramidal
panicle; flowers blue; tropical and subtropical.
22. Coleus armticus Benth. (Fig. 23a) 'Wild Oregano' or 'Indian Borage';
origin-Malaya and Indonesia; fleshy perennial herb; stem weak, obturely
4-angled, pubescent; ,leaves succulent, ovate-elliptic, margins finely
serrate, light green, puberulous, petioles slender; tropical and
subtropical.
23. C. arovlticus Benth. var.variegata (Fig. 24a) 'Variegated Borage',
perennial herb; stem fleshy, puberulous; leaves showy, variegated, thick,
margin crenate, obovate, finely tomentose; petioles greenish white;
tropical and subtropical.
24. C. bl-i Benth. var verschaffeltii (Fig. 25a) 'Painted nettle'; origin-
Java; perennial herb; stem square, half hardy, purple; leaves ovate,
margin toothed, showy, blade shows crimson coloured patches, purple in
centre, border narrow green; petiole purplish-white; inflorescence a
branched spike; flowers pale blue; tropical.
25. C. forskohlii Briq. (Fig. 26a) Perennial herb; stem obtusely 4-angled,
woody at base, pubescent; leaves large, thick, ovate-oblong, hispidly
tomentose on both sides, margins crenate; tropical and subtropical.
26. C. laciniatus Benth. (Fig. 27a) Perennial herb; stem fleshy; leaves soft,
torn or slashed into narrow lobes, attractively coloured; inflorescence a
branched lax spike; flowers light purple; tropical.
27. C. parviflorus Benth. (Fig. 28a) 'Country potato', perennial herb; stem
tetragonous, fleshy, produces tubers at the base; leaves long petioled,
membraneous, chartaceous, margin crenate, rarely pubescent, grassy green;
tropical.
28. C. rehneltianus Berger (Fig. 29a) 'Red Trailing Queen'; perennial herb;
robust, plant with half-hardy, quadrangular, purple stem; leaves broad-
ovate, margins slightly lobed, blade deep purple coloured with carmine in
the centre and a broad border of dullgreen; tropical.
29. C. zeylanlcus (Benth.) Cramer (Fig. 30a) An aromatic, perennial herb;
stem obscurely tetragonous, pubescent, light green in tender parts,
slightly brownish when old; leaves large, thin, soft, velvetty, rounded,
puberulous, yellowish green; tropical and subtropical.
Fig. 23a Fig. 24a
Fig . 22a
F i g . 26a Fig. 27a
30. Anisochilu carnosus (L.f.) Wall. (Fig. 31a) A gregarious annual
subshrub; stem stout, bluntly 4-angled, glabrous, often tinged with red;
leaves rather fleshy, ovate-cordate, pubescent, margin serrate,
deciduous; inflorescence a strobiloid head like spike; flowers
palepurple; tropical.
31. A. eriocephalu Benth. (Fig. 32a) Annual herb; stem and branches
puberulous; leaves ovate-orbicular, crenate, dark green; inflorescence a
dense cylindrical spike; flowers lavender blue; tropical.
32. Hyptis capitata Jacq. (Fig. 33a) Annual subshrub; stem half-hardy,
distinctly tetrangular, sparingly pubescent; leaves large, ovate-oblong,
margin crenate-serrate, coarse, dark green; inflorescence a capitate
head, axillary; flowers minute white; tropical and subtropical.
33. H. suaveolens (L . ) Poit. (Fig. 34a) Origin-Tropical America and West
Indies; aromatic sweet smelling annual herb; stem tetragonal, roughly
hairy; leaves ovate-cordate, denticulate, strigose; inflorescence a
verticillaster of stalked cymes; flowers small, blue; tropical.
34. Pogostemon benghalensis (Burm.f.1 Kuntze (Fig. 35a) Perennial herb; stem
and branches subquadrangular, covered by grey tomentose; leaves ovate-
oblong, doubly serrate, puberulous; inflorescence an erect terminal
panicle formed of many short spikes; flowers pink-purple; tropical and
subtropical.
35. P. heyneanus Benth.(Fig. 36a) Perennial undershrub; stem and branches
glabrous, nodes swollen, obliquely 4-angled, brownish green; leaves
large, ovate, sparingly puberulous, greyish green; tropical and
subtropical.
4 Fig. 29a
Ffg. 33a
Fig. 31a
Fig. 37a
36. P. purpurascens Dalz. (Fig. 37a) Perennial herb; stem half-hardy,
branchlets hispid; leaves in unequal pairs, ovate-acuminate, inciso-
serrate, pubescent; inflorescence a compact panicle, formed of flattened
spikes; flowers purplish-white; tropical and subtropical.
37. P. vestitus Benth (Fig. 38a) Perennial herb; stem densely wooly; leaves
ovate-obtuse, serrate, pubescent; inflorescence a unilateral panicle,
bracts foliaceous; flowers purple, small; tropical.
38. Eusteralis quadrifolia (Benth.) Panigrahi (Fig. 39a) An annual aromatic
woody herb; leaves in whorls of four, narrowly oblong-obtuse, clothed by
dense greyish tomentose; inflorescence a long dense terminal spike;
flowers rosy pink, minute; tropical.
39. Hentha amensis L. (Fig. 40a) 'Japaneese mint' or 'Field mint'; origin
Japaneese Archipelago; perennial aromatic herb with running leafy
stolons; stem square, branches ascending; leaves broadly oblong,
puberulent, sharply serrate; temperate and subtropical.
40. M. rotundifolia (L.) Huds. (Fig. 41a) 'Round-leaved mint'; origin-
Temperate Asia; perennial herb with running root stock; stem slender
without much branches, glabrous; leaves nearly round, crenate, obtuse,
dark green; temperate and subtropical.
41. H. spicata (L.) Huds. (Fig. 42a) 'Spear mint'; origin-Temperate Europe
and Asia; perennial herb; rootstocks spreading; stem erect; leaves
oblong-lanceolate, serrate, much wrinkled, chartaceous, finely pubescent
on the veins; temperate and subtropical.
42. Salvia coccinea Juss. (Fig. 43a) 'Red sage' or 'Texas sage'; origin-
Central and Tropical Aierica; perennial undershrub; stem sparsely hairy;
leaves subcordiform, puberulous, crenate-serrate, gray-green;
inflorescence a lax raceme; flowers deep red; tropical and subtropical.
43. S. leucantha Cav. (Fig. 44a) 'Mexican bush sage'; origin-Central Mexico;
perennial wooly undershrub; stem and branches tomentose; leaves, narrow,
oblong-lanceolate, crenate, blade white lanate beneath; inflorescence a
long slender spike, floral whorls remote at the base; flowers purple,
covered by downy felt; subtropical.
44. S. plebeia R.Br. (Fig. 45a) A roughly pubescent annual subshrub; stem
stout, fastigiately branched, obtusely 4-angled and grooved; leaves
oblong-lanceolate, crenate-serrate, rugose; inflorescence a panicle of
compact racemes; flowers small, blue; tropical.
45. S. splendens K&- awl. (Fig. 46a) 'Scarlet sage'; origin - Brazil;
perennial undershrub; stem half hardy, branches glabrous; leaves ovate-
acuminate, crenate-serrate; inflorescence a terminal spike like raceme;
flowers showy, scarlet-red; tropical.
46. Adscaeles indica (L.)Kuntze (Fig. 47a) Perennial under shrub; stem
acutely tetragonous, covered with greyish tomentose, branches softly
hairy; leaves obovate, chartaceous, crenate, pubescent; inflorescence,
terminal or axillary dense spikes formed of sessile cymes; flower
purplish - white; tropical.
47. A lalabarica R.Br. (Fig. 48a) Perennial undershrub; stem obtusely
tetragonous, pubescent; leaves oblong-lanceolate, sub-coriaceous;
crenate-serrate, blade pale green above and white below, softly wooly;
inflorescence a dense spike; flowers pale purple, pubescent; tropical.
F i g . 44a
Fig* 3%.
Fig. 42a
Fig. 45a F i g . 46a
48. Leuas aspera Spreng. (Fig. 49a) Coarse annual herb; stem square, grooved
on all sides, rugose; leaves linear-oblong, toothed, coarse hairy;
inflorescence a verticillaster; flowers white, sessile; tropical.
49. L. cephalotes Spreng. (Fig. 50a) Annual herb; stem sparsely pubescent,
tetragonous; leaves linear-oblong, toothed, roughly hairy; inflorescence
a verticillaster, slightly pendant; flowers white; tropical.
50. L. linifolia Spreng. (Fig. 51a) Annual herb; stem 4-angled, pubescent,
branched below; leaves linear-lanceolate, narrow, almost entire, covered
by appressed hairs; inflorescence a verticillaster; flowers white,
shortly stalked, finely pubescent; tropical.
51. L. stricta Benth. Annual herb; stem upright, stout; leaves narrow,
coarse, elliptic-lanceolate; densely hairy; inflorescence a
verticillaster; flowers white; tropical.
52. L. vestita Benth. (Fig. 53a) Annual herb; stem quadrangular, tawny
villous with brownish felt; leaves lanceolate, crenate-serrate,
pubescent; inflorescence a verticillaster; flowers white, bearded;
subtropical.
53. Leo~tis nepetifolia ( L . ) Ait.f. (Fig. 54a) Origin-Tropical America;
perennial glabrous undershrub; stem 4-angled, stout, erect; leaves ovate-
elliptic, soft, pubescent, crenate-serrate; inflorescence a
verticillaster formed of gorgeous many flowered whorls; flowers showy
orange- red with hooded downy corolla and spinescent calyx; tro.pica1.
54. Teucrium plectranthoides Gamble (Fig. 55a) Perennial undershrub; stem
branches robust, quadrangular, glabrous; leaves long, greyish green,
finely serrate; inflorescence a spreading panicle of open racemes;
flowers yellowish white; tropical.
Fig. 50a
Fig. 48a Fig. 49a
47a-Habit of Anisomeles indica (L.) Kuntze, 48a-A.malabarica - R . B r . , 49a-Lcucas aspera Spreng., 50a-L. cephalotes Spreng., 51a-L.linifo1ia Spreng., 52a-L.stricta Benth. 53a-%.vestita Benth., 54a-Leonotis nepetifolia (L.) Ait.f., 55a-Teucrium plectranthoizes Gamble.
METHODS
A. Cytological studies
1. Mitotic squash experiments
Somatic chromosome studies have been conducted on fifty four members
of South Indian Lamiaceae with the help of improved cytotechniques
(Sharma and Sharma, 1980). Squash experiments were carried out on
root tip meristem at mitotic metaphase stage. Young healthy root
tips are collected from the vegetative cuttings planted in the
experimental botanical garden, at the periods showing peak mitotic
frequency, i.e., 9.00 A.M. to 10.30 A.M. In plant cutting with a
lesser rooting ability, seeds are germinated in petridishes lined
with moist filter paper under laboratory conditions. The root tips
are then washed thoroughly in distilled water and pretreated with
different pretreating chemicals. Saturated aqueous paradichloro
benzene solution with a trace of aesculine is found to be most
suitable for the members of Lamiaceae. The root tips are initially
chilled at 0 - 5 O ~ for 4-6 minutes and then kept at 12-14O~ for 1-3
hours for obtaining the best results.
The pretreated root tips are then washed thoroughly with distilled
water and fixed in 1:3 acetic acid - ethyl alcohol mixture (Carnoy,
1886) overnight, followed by 3-7 minutes treatment in 45% acetic
acid and stained with 2% aceto-orcein - 1 N HC1 mixture (9:l) for 2
112 to 3 112 hours. While staining, the vial is initially warmed
over a flame for effective results. The stained root tips are then
squashed in 45% acetic acid and the preparation was then temporarily
sealed.
To remove the excess oil deposits, rather common in Lamiaceae, it
has been found preferable to add a little saponin along with the
pretreatment mixture and shake thoroughly.
2. Meiotic smear experiments
pollen mother cell (PMC) analysis were carried out on those members
which bloomed frequently in the experimental garden. For this flower
buds are fixed in modified Carnoy's fluid (1 acetic acid : 3
chloroform : 6 ethyl alcohol) for 1-2 days and stored in 70% ethyl
alcohol. Young anthers are smeared with 2% aceto-carmine stain to
obtain PMC's showing divisional stages.
For better results, the fixing mixture is replaced by fresh Carnoy's
fluid one or two times, which helpes to remove the excess cell
inclusions.
B. Detailed Karyaorphometrical analysis
1. Drawings - and Wotoricrographs
Karyotype drawings are made with the help of a Leitz-BIOMED tracing
&vice with a magnification of X 2400 (approximately). The
photomicrographs are taken using 125 ASA 35mm ORWO film in a Leitz
photographic attachment and suitably enlarged. The idiograms are
prepared after comparing atleast five well spread metaphase plates.
2. Karyomorphological studies
The total length as well as the short arm length of all the
chromosomes of the fifty four cytotypes are measured accurately
using micrometers. In all the karyotypes, ratio of the short arm to
the total length of the chromosome in percentage, F% (Forma
percentage or Centromeric index) is determined after Krikorian, et
al. (1983). On the basis of F% the nature of primary constriction in
the chromosomes are classified as follows:
The disparity index (DI) of chromosomes in a karyotype is
calculated after Mohanty, et al. (1991), by the formula:
F%
50
49.9 - 37.6 37.5 - 25.1
25
24.9 - 18.76
18.75 - 12.6
12.5
12.4 - 6.26
6.25 - 1 1
DI = Longest chromosome - Shortest chromosome x 100 Longest chromosome + Shortest chromosome
Nature of Primary Constriction
Median
Nearly median
Nearly submedian
Submedian
Nearly submedian
Nearly subterminal
Subterminal
Nearly subterminal
Extremely subterminal
Terminal
-
The variation coefficient among chromosome complements (VC) is
determined after Verma (1980) as follows:
VC = Standard deviation X 100 Mean length of chromosomes
The total forma percentage or the mean centromeric index value (TFX)
is calculated in each taxa after Huziwara (1962), by the formula :
TF% = Total sum of short arm lenth X 100 - - - - - - - - Total sum of chromosome lenth
For the calculation of polyploidy, the base numbers given in
Darlington and Wylie (1955), Morton (1956a, 1962), as well as Love
and Love (1961b), are usually followed. In those cases where the
basic numbers are not mentioned in these references, the latest
literature is consulted.
In the present investigation, fifty four members of South Indian Lamiaceae including
twelve cytotypes in the genus Ocimum L., one in Acrocephalus Benth., six in
Orthosiphon Benth., two in Plectranthus L' Herit., eight in Coleus Lour., two in
Anisochilus Wall., two in Hyptis Jacq., four in Pogostenon Desf., one in Eusteralis
Rafin., three in Mentha L., four in Salvia L., two in Anisomeles R.Br., five in
Leucas R.Br., one in Leonotis R.Br. and one in Teucrium L. were analysed. The
normal somatic chromosome number ranges from 2n = 16+1-OB to 96. However,
numerical variations have been recorded in many species. The ploidy level exhibited
by different taxa range from diploidy to octaploidy. The chromosome pair with
secondary constriction is found to range between one and three. The karyotypes are
characterized by the presence of comparatively small chromosomes, ranging from 4.2 p
to 1.0 p in length. The chromosomes in each karyotype decrease in size progressively
and bear nearly median to nearly submedian primary constriction.
The general description of the common chromosome types is given below, followed by a
karyotype description for the members investigated.
Type A : Comparatively long chromosome with two constrictions, one median and the
other nearly sibmedian in position (4.2 to 1.4 p).
Type B : Comparatively long chromosome with nearly median to nearly submedian
primary constriction and a satellite at the distal end of shorter arm,
joined by a SAT thread (2.8 to 2.2 p) .
Type C : Relatively long chromosomes (4.2 - 2.0~) with nealy median to nearly
submedian primary constriction.
Type D : Medium to short chromosomes (<2.0p) with nearly median to nearly
submedian primary constriction.
Diagrammatic representation of the different chromosome types observed in the South Indian taxa of Lamiaceae is shown below.
L-__r . 'I'y A 'I 'y IN: II
Diagrammatic representation of different chromosome
types found in the taxa investigated in Laoiaceae
Chromosome count on pollen mother cell
Normal somatic chromosome number
Somatic variation numbers
Chromosome pairs with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
n = 11 (Fig. 2j)
2n = 22 (Fig. 2b, 2-2)
2n = 24 (Fig. 2e)
1
3 . 2 ~ to 1 . 8 ~
2.31~
50.
25.4 p
28.00
14.93
40.08
Table 2 --
Detailed karyotype analysis (Pig. 2d)
Chromo- some type
A
C
C
C
C
D
No. of pairs
1
4
1
2
2
1
Total length in AI
3.2
2.4
2.4
2.2
2.0
1.8 --
Nature of primary constriction
-
Nearly median
Nearly submedian
Nearly median
Remarks
Chromosome pair with secondary constriction
-
-
- -
-
Short arm length in AI
0.6 0.8
1.0
0.8
0.8
0.8
0.8
' FX
18.75 25.00
41.67
33.33
36.36
40.00
44.44 --
Chromosome count on pollen mother cell : n = 36 (Fig 3 j )
Normal somatic chromosome number : 2n = 72 (Figs. 3b, 3c)
Somatic variation numbers : 2n=18, 40 and 48 (Figs. 3e,3f ,3g)
Chromosome pair with secondary constriction : 2
Range of chromosome length : 4.2 p to 1.4 p
Average chromosome length : 2 . 1 5 ~
Total chromatin length : 154.8 p
Chromatin length of basic complement : 25.8 p
Disparity index (DI) : 50
Variation coefficient (VC) : 28.28
TF value (%) : 43.04
Table 3 --
Detailed karyotype analysis (Pig. 3d)
Chromo- some type
A
A
C
C
C
C
C
C
C
D
D
D
Short arm length in p
1 .0 1.2
0.8 1.0
1.4
1.2
1 . 0
1 .0
1.0
0 .8
0.8
0 . 8
0.6
0.6
No. of pairs
1
1
1
1
2
2
4
3
5
7
7
2
F%
23.81 28.57
23.53 29 .41
43.75
40.00
35.71
38.46
41.67
36.36
40.00
44.44
37.50
42.86
Total length in u
4.2
3.4
3.2
3.0
2.8
2.6
2.4
2.2
2.0
1.8
1 .6
1.4
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
Nearly submedian
Nearly median
Nearly submedian
Nearly median
Remarks
Chromosome pair with secondary constriction
-
- -
-
-
-
-
-
-
-
A\ p?
O c h u m basilicum L.var. glabratum Benth. (2n=4x=48=A4C12D32)
Chromosome count on pollen mother cell : n = 24 (Fig. 4j)
Normal somatic chromosome number : 2n = 48 (Figs. 4b,4c)
Chromosome pair with secondary constriction : 2
Range of chromosome length : 2.2 p to 1.2 p
Average chromosome length : 1.73 p
Total chromatin length : 83.2 p
Chromatin length of basic complement
Disparity index (DI) : 29.41
Variation coefficient (VC) : 16.57 s ,
TF value (%I : 40.57.'-. . ,
Table 4 --
Detailed karyotype analysis (Fig.4d)
Remarks
Chromosome pair with secondary constriction
-
-
-
-
-
-
-
Chromo- some type
A
A
C
C
D
D
D
D
D
F%
18.18 27.27
18.18 36.36
45.45
40.00
33.33
44.44
37.50
42.86
33.33
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
Nearly submedian
Nearly median
Nearly submedian
No. of pairs
1
1
1
5
2
2
6
5
1
Total length in p
2.2
2.2
2.2
2.0
1.8
1.8
1.6
1.4
1.2
Short arm length in p
0.4 0.6
0.4 0.8
1.0
0.8
0.6
0.8
0.6
0.6
0.4
Fig. 2c
Fig. 3g
Fig. 3c
Fig. 3f
# Fig. 4c
Fig. 2c, 2d - Karyotype and idiogram of -- Ocimum adscendens Willd. (2n=22)x2400 approx., 3c, 3d, 3f, 3g - - 0. americanum L.: 3c; 3d Karyotype and idiogran ( 2 ~ 7 2 ) x 2400 amrox.. 3f. 3~ somatic variants with 211-40 and 2n=48 , - respectively; 4c, ii - bryotype and idiograrn of - 0 basilicum L.var.&abratun Benth. (2n=48) x 2400 approx.
-
Fig. 2b .Fig. 2e Fig. 2j
- Fig. 3b Fig. 3 j
- -
Fig. 4b Fig. 4j Pig. 5b
Scale: r 8 wJ
Figs. 2b, 2e, 2 j - Oci mum adscendens Willd. : 2b mitotic metaphase (2n=22), - 212 - somatic variant (2n=24), 2j - meiotic anaphase I (n=ll); 3b, 3e, 33 - 2. americanum L.: 3b - mitotic metaphase ( 2 ~ 7 2 1 , 3e - somatic variant (2nX18), 3j - meiotic anaphase I (abnormal seggregation); 4b, 4j - 0. basilicum L. var. - labracum Benth.: 4b - mitotic metaphase (2n=48), 4j - meiotic metaphase I
fn=24); 5b - - 0 . bnsilicum L. var. pilosum Benth. - mitotic metaphase (2n=48).
Oc-um basilium L.var. pilosum Benth. (2n=4~=48=A4CZOD24)
Chromosome count on pollen mother cell
Normal somatic chromosome number
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
n = 24 (Fig. 5j)
2n = 48 (figs. 5b, 5c)
2
3.2 p to 1.4 p
1.96 1.1
94 1.1
23.5 p
39.13
19.76
41.14
Table 5 --
Detailed karyotype analysis (Pig. M)
Remarks
Chromosome pair with secondary constriction
-
-
-
-
-
-
F%
25.00 31.25
23.08 30.77
41.67
36.36
40.00
44.44
37.50
42.86
Chromo- some type
A
A
C
C
C
D
D
D
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
Nearly submedian
Nearly median
Total length in p
3.2
2.6
2.4
2.2
2.0
1.8
1.6
1.4
No. of pairs
1
I
1
4
5
5
6
1
Short arm length in p
0.8 1.0
0.6 0.8
1.0
0.8
- 0.8
0.8
0.6
0.6
O c i n n basillcum L. var . purpurascens Benth. (211=4~=48=A4C28D16)
Chromosome count on pollen mother cell : n = 24 (Fig. 6j)
Normal somatic chromosome number : 2n = 48 (Figs. 6b, 6c)
Chromosome pairs with secondary constriction : 2
Range of chromosome length : 2.8 p to 1.6 1.1
Average chromosome length
Total chromatin length
Chromatin length of basic complement : 2 5 . 1 ~
Disparity index (DI) : 27.27
Variation coefficient (VC) : 15.13
TF value (%) : 42.79%
Table 6 -- Detailed karyotype analysis (Pig. 6d)
Chromo- some type
A
A
C
C
C
C
C
D
D
No. of pairs
1
1
1
3
4
1
5
6
2
Total length in 1.1
2.8
2.6
2.6
2.4
2.2
2.2
2.0
1.8
1.6
Short arm length in 0
0.6 0.8
0.6 0.8
1.0
1.0
1.0
0.8
0.8
0.8
0.6
F%
21.43 28.57
23.08 30.77
38.46
41.67
45.45
36.36
40.00
44.44
37.50
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
Nearly submedian
Remarks
Chromosome pair with secondary constriction
-
- -
- -
- -
Ocimum basilicum L.var. thyrsiflorum Benth. (2n=4~=48=A4C22D22)
Chromosome count on pollen mother cell : n = 24 (Fig. 7 j )
Normal somatic chromosome number : 2n = 48 (Figs. 7 b , 7c)
Chromosome pair with secondary constriction : 2
Range of chromosome length : 3.4 fl to 1.4 f l
Average chromosome length : 2.02 /.I
Total chromatin length : 96.8 p
Chromatin length of basic complement : 24.2 p
Disparity index (DI) : 41.67
Variation coefficient (VC) : 21.94
TF value (%) : 40.39
Table 7 -- Detailed Laryotype analysis (Pig. 7d)
Chromo- some type
A
A
C
C
C
C
D
D
D
D
No. of pairs
1
1
1
2
2
6
2
3
5
1
Total length in p
3.4
2.8
2.6
2.4
2.2
2.0
1.8
1.8
1.6
1.4
Short arm length in p
0.8 1.0
0.6 0.8
1.0
0.8
1.0
0 .8
0.6
0.8
0.6
0.6 --
F%
23.53 29.41
21.43 28.57
38.46
33.33
45.45
40.00
33.33
44.44
37.50
42.86
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
Nearly submedian
Nearly median
Nearly submedian
Nearly median
Remarks
Chromosome pair with secondary constriction
-
-
- - - -
- -
Fig. 5c Fig. 7c
Fig. ,6c
Fig. 5c, 5d - Karyotype and idiogram of Ocimum basilicum L.var.pilosum Benth. i2n=48) x 2400 approx. 6c,bd - Karyotype and idiogram of 0 . basilicum L. var.purpurascen Benth. ( 2 ~ 4 8 ) x 2400 approx.; 7c,7d - ~aryoty~e-and idiogram of 0 . basilicum L var. thyrsiflorum Benth ( 2 ~ 4 8 ) x 2400 approx. - .- -
Chromosome count on pollen mother cell : n = 20 (Fig. 8j)
Normal somatic chromosome number : 2n = 40 (Figs. 8b, 8c)
Somatic variation numbers : 2n=34 and 42 (Figs. 8 e , 8 f )
Chromosome pairs with secondry constriction : 2
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement : 2 0 . 1 ~
Disparity index (DI) : 27.27
Variation coefficient (VC) : 17.65
TF value (%) : 40.66
Table 8 --
Detailed karyotype analysis (Pig. 8d)
Remarks
Chromosome pair with secondary constriction
Chromosome pair with a satellite
-
- -
-
-
-
Chromo- some type
A
B
C
C
C
D
D
D
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
Nearly submedian
Nearly median
Nearly submedian
No. of pairs
1
1
2
2
5
2
3
4
Total length in p -
2.8
2.8
2.4
2 .2
2.0
1 . 8
1 .8
1.6
Short arm length in p
0 . 6 0 .8
0 . 8 0 .8
1 .0
0 . 8
0 .8
0 . 6
0 .8
0 . 6
F%
21.43 28.57
28.57 28.57
41.67
36.36
40.00
33.33
44.44
37.50
- -- - ---
Fig. 5j Fig. 6b Fig. 6 j
-- - -
Fig. 7b Fig. 7j Fig. 8b
Fig. 8e Fig. 8j Fig. 8m
Scale: I.----r BP' - --!
Fig. 5j - Ocimum basilicum L. var. pilosum Benth.- meiotic metaphase I ( ~ 2 4 ) ; 6b, 6 j - - 0 . basilicum L. var. purpurascens Benth.: 6b - mitotic metaphase (2n=48), 6j-meiotic metaphase I (n=24); 7b, 7j - - 0. basilicum L. var. thyrsiflorum Benth.: 7b - mitotic metaphase (2n=48), 7j - meiotic anaphase I (n=24); 8b, 8e, 8j, 8m - - 0. gratissimum L.: 8b - mitotic metaphase (2nz40), 8e - somatic variant (2n = 34), 8j - meiotic metaphase I (abnormal with 14 bivalents and 12 univalents), 8m - normal tetrad.
Ocimum gratissimum L.var. suavis H0ok.f. (2n=4~=48=A2B2C20D24)
Chromosome count on pollen mother cell : n = 24 (Fig 9j)
Normal somatic chromosome number
Somatic variation numbers
: 2n = 48 (Fig. 9b, 9c)
: 2 n ~ 3 6 , 38and42 (Figs. 9e,9f ,9g)
Chromosome pair with secondary constriction : 2
Range of chromosome length : 2.6 p to 1.4 p
Average chromosome length : 1.9 p
Total chromatin length : 91.2 p
Chromatin length of basic complement : 22.8 p
Disparity index (DI) : 30.00
Variation coefficient (VC) : 15.19
TF value (%) : 42.64
Table 9 -- Detailed karyotype analysis (Pig. 9d)
Chromo- some type
A
B
C
C
C
D
D
D
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
Remarks
Chromosome pair with secondary constriction
Chromosome pair with a satellite
- -
-
-
-
-
No. of pairs
1
1
1
2
7
6
4
2
Short arm length in p
0.6 0.8
0.6 0.6
1.0
1.0
0.8
0.8
0.6
0.6
Total length in p
2.6
2.2
2.4
2.2
2.0
1.8
1.6
1.4
F%
23.08 30.77
27.27 27.27
41.67
45.45
40.00
44.44
37.50
42.86
Fig. 8c
Fig. 9e
Fig. 8f
Fig. 9f
Fig. 9c
Fig. 9g
Fig. 8c,8d,8f - Ocimum gratissimum L.: 8c, 8d - Karyotype and idiogram ( 2 ~ 4 0 ) x 2400 approx., 8f - somatic variant (2n=42); 9c, 9d, 9e, 9f. 9g - 0. gratissirnum L.var.suavis H0ok.f. : 9c,9d - Karyotype and idiogram (2n=48) x 2400 approx., 9e,9f,9g - somatic variants with 2n=36, 2n=38 and 2n=42 respectively.
Ocimm tenuiflonm L. f . cv. green (2n=4x=32=A4C4D24)
Chromosome count on pollen mother cell
Normal somatic chromosome number
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
n = 16 (Figs. 10j)
2n = 32 (Figs. lob, 10c)
2
2.0 @ to 1.2 p
1.65 ,u
52.8 p
13.2 p
25.00
16.87
43.18
Table 10 --
Detailed Karyotype analysis (Fig. 10d)
Remarks
Chromosome pair with secondary constriction.
- -
- -
-
I I I
Nature of primary constriction
-
Nearly median
Nearly submedian
Nearly median
Nearly submedian
Short arm length in p
0.6 0.6
0.8
0.8
0.6
0.6
0.4
Total length in ,u
2.0
2.0
1.8
1.6
1.4
1.2
Chromo- some type
A
C
D
D
D
D
F%
30.00 30.00
40.00
44.44
37.50
42.86
33.33
No. of pairs
2
2
4
2
4
2
Ocinn tenuiflonm L.f. cv. purple (2n=4x=32=A4D28)
Chromosome count on pollen mother cell
Normal somatic chromosome number
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
n = 16 (Fig. llj)
2n = 32 (Figs. llb, llc)
2
1.8 ,u to 1.0 ,u
1.49 p
47.6 p
11.9 u
28.57
18.36
35.93
Table 11 --
Detailed Karyotype analysis (Fig. lld)
Remarks
Chromosome pair with secondary constriction.
- -
-
- -
Nature of primary constriction
-
-
Nearly submedian
D 1 2 1 1.0 I 0.4 1 40.00 I Nearly median
F%
22.22 33.33
25.00 25.00
33.33
37.50
28.57
33.33
Short arm length in p
0.4 0.6
0.4 0.4
0.6
0.6
0.4
0.4
Total length in p
1.8
1.6
1.8
1.6
1.4
1.2
Chromo- some type
A
A
D
D
D
D
No. of pairs
1
1
4
2
4
2
-- -- - - - - .- -
Fig. 8n Fig. 80 Fig. 8 p
- -. -- Fig. 9b Fig. 9 j Fig. lob
Fig. 1 O j - -- Fig. l l b
Scale: - 73 I C ~
Fig. l l j
Figs. an , 80, 8p - Ocimum gratissimum L. showing polyspory: 8n - pentad, 80 - hexad, 8p - octad; 9b, 9j - - 0 . - gratissimum L.var. suavis Hook. f.: 9b - mitotic metaphase (2n=48), 9 j - meiotic anaphase I (abnormal seggregation); lob, 10 j - 0 . tenuiflorum L . f . cv. p e e n : lob - mitotic metaphase (2n=32), 10j. - meioti; meta~hase I (n=16) ; I l b , 11 j - 0 . tenuiflorum ~ . f . cv. purple: l l b - mitotic metaphase I (2n=32), 11 j - diakiyesis (n=16).
39
Ocimm tenuiflonm L.f. cv. purple-green (2n=4x=36=A4D32)
Chromosome count on pollen mother cell
Normal somatic chromosome number
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
T F value (%)
n = 18 (Fig. 12j )
2n = 36 (Figs. 12b, 12-2)
2
1 . 8 p to 1 . 0 ,u
1 . 3 8 p
Table 12 --
Detailed Karyotype analysis (Pig. 12d)
Chromo- some type
A
D
D
D
D
D
I l l I I I
No. of pairs
2
2
2
2
2
2
Total length in p
1 . 8
1 .8
1 .6
1 . 4
1 . 2
1 . 0
Short arm length in p
0 . 4 0 . 6
0 . 6
0 . 6
0 . 6
0 . 4
0 . 4
F%
22.22 33 .33
33.33
37.50
42.85
33.33
40.00
Nature of primary constriction
-
Nearly submedian
Nearly median
Nearly submedian
Nearly median
Remarks
Chromosome with secondary constriction.
- - - -
-
Chromosome count on pollen mother cell
Normal somatic chromosome number
Chromosome pairs with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value ( X I
n = 18 (Fig. 1 3 j )
2n = 36 (Fig. 13b, 13c)
2
Table 13 --
Detailed karyotype analysis (Pig. 13d)
Remarks
Chromosome pair with secondary constriction
-
-
-
-
- -
Chromo- some type
A
A
C
D
D
D
D
D -
F%
20.00 30.00
22.22 33.33
40.00
44.44
37.50
42.86
33.33
40.00
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
Nearly submedian
Nearly median
No. of pairs
1
1
2
2
3
6
2
1
Total length in p
2.0
1.8
2.0
1 .8
1.6
1.4
1.2
1.0
Short arm length in p
0.4 0.6
0.4 0.6
0.8
0.8
0.6
0.6
0 .4
0.4
Fig. 1oc Fig. llc
Fig. 12c Fig. 13c
Fig. 10d J8 88, ,#I 11, 11 8 l 1 1 8 ) 8 ) 4 1 I1 11 Il I 1 18,
Fig. lld jfi dd, J1 4; 18 11 18 11 I1 I t I 8 81 81 88 18,
Fig. 10c, 10d - Karyotype and idiogram of -- Ocinun tenuiflorum L.f. cv ereen (2n=32) x 2400 approx., llc, lld - Karyotype and idiogram of 2. tenuiflorm L.f. cv.purple f2n=32) x 2400 approx., 12c, 12d - karyotype and idiogram of 0 tenuiflorum L.f cv purple-green (211-36) x 2400 approx.; 13c, 13d - - Karyotype and idiogran of 2. tenuiflorum L.f. var. hirsuta (2n=36) x 2400 approx
Acrocephalus capitatus Benth. (2n=Zx=l8=AZD16)
Normal somatic chromosome number
Chromosome pair w*.th secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
2n = 18 (Figs. 14b, 14c)
1
1 . 6 ~ to 1.0~
1.36~
24.b
12.w
23.08
13.47
40.62
Table 14 -- Detailed karyotype analysis (Fig. 14d)
Remarks
Chromosome pair with secondary constriction
-
- - -
Chromo- some type
A
D
D
D
D
F%
25.00 25.00
37.50
42.86
33.33
40.00
Nature of primary constriction
-
Nearly median
Nearly submedian
Nearly median
No. of pairs
1
1
4
2
1
Total length in p
1.6
1.6
1.4
1.2
1 .O
Short arm length in p
0.4 0.4
0.6
0.6
0.4
0.4
Orthosiphon glabratus Benth. (2n=Zx=26=AZC24)
Chromosome count on pollen mother cell
Normal somatic chromosome number
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
n = 13 (Fig. 15j)
2n = 26 (Figs. 15b, 15c)
1
3.6 p to 2.2 p
2.74 p
71.2 p
35.6 p
24.14
13.24
39.93
Table 15 --
Detailed Karyotype analysis (Pig. 15d)
Remarks
Chromosome pair with secondary constriction.
-
- -
Chromo- some type
A
C
C
C
F%
22.22 33.33
40.00
38.46
36.36
Nature of primary constriction
-
Nearly median
Nearly submedian
No. of pairs
1
4
6
2
Total length in pl
3.6
3.0
2.6
2.2
Short arm length in &I
0.8 1.2
1.2
1.0
0.8
Orthosiphon glabratus var. parviflorus Benth. (2n=2~=26=A2ClZD12)
Chromosome count on pollen mother cell
Normal somatic chromosome number
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
n = 13 (Fig. 16j)
2n = 26 (Figs. 16b, 16c)
1
2.8 ,u to 1.4 p
2.06 ,u
53.6 ,u
26.8 ,u
33.33
20.64
41.22
Table '16 --
Detailed Karyotype analysis (Pig. 16d)
Remarks
Chromosome pair with secondary constriction.
- - - -
Chromo- some type
A
C
C
D
D
F%
21.43 28.57
38.46
36.36
44.44
42.86
Nature of primary constriction
-
Nearly median
Nearly submedian
Nearly median
No. of pairs
1
2
4
4
2
Total length in ,u
2.8
2.6
2.2
1.8
1.4
Short arm length in p
0.6 0.8
1.0
0.8
0.8
0.6
-- --- Fig. 12b Pig. 12; Fig. 13b
--- - - .- - -. - Fig. 13j Fig. 14b Fig. 15b
-- - - - --
Fig. 15j Fig. 16b Fig. 16j
Scale: 811h
Figs. 12b, 12j - Ocimum tenuiflorum L.f. cv.purple-green: 12b - mitotic metaphase (2n=36), 12j-meiotic anaphase I (n=18); 13b, 13j - - 0. tenuiflorum L . f . var. hirsuta H0ok.f.: 13b - mitotic netapllase (2n=36), 13j - meiotic metaphase 1: (n=18); 14b - Acrocephalus capitatus Benth. - mitotic metaphase (2n=,18); lSb , 1 5 j - Orthosiphon glabratus Benth.: 15b - mitotic metaphase (2n=26), 15j - meiotic anaphase 1 (~13); 16b, 1 6 j - 0. glabratus var. parviflorus Benth. : 16b - mitotic metaphase (2~261, 16 j-. meiotic metaph- 'ase I (n=13)
Orthosiphon grandiflorus Bold cv. lilac (Zn=4~=28=A2ClZD14)
Chromosome count on pollen mother cell
Normal somatic chromosome number
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (X)
n = 14 (Fig. 17j)
2n = 28 (Figs. 17b, 17c)
1
2.8 ,u to 1.4 p
2.04 p
57.2 p
14.3 p
33.33
21.66
38.6
Table 17 -- Detailed Karyotype analysis (Pig. 17d)
Remarks
Chromosome pair with secondary constriction.
- - -
- -
Chromo- some type
A
C
C
C
D
D
D I I
FX
21.43 35.71
38.46
36.36
40.00
33.33
37.50
No.of pairs
1
3
2
1
3
3
0.6 42.86 I Nearly median I -
Nature of primary constriction
-
Nearly median
Nearly submedian
Nearly median
Nearly submedian
Total length in p
2.8
2.6
2.2
2.0
1.8
1.6
Shortarm length in p
0.6 1.0
1.0
0.8
0.8
0.6
0.6
Orthosiphon grandiflorus Bold cv. white (2n=4~=28=AZC18D8)
Chromosome count on pollen mother cell
Normal somatic chromosome number
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
n = 14 (Fig. 181)
2n = 28 (Figs. lab, 18~)
1
3.2 p to 1.6 ,u
2.2 cl
61.6 p
15.4 p
33.33
22.00
42.1
Table I8 -- Detailed Karyotype analysis (Fig. 18d)
Chromo- some type
A
C
C
C
C
C
D
I I I I I I
No. of pairs
1
1
2
2
2
2
4
Total length in p
3.2
2.8
2.6
2.4
2.2
2.0
1.6
Short arm length in p
0.6 1.0
1.2
1.2
1.0
1.0
0.8
0.6
FX
18.75 31.25
42.86
46.15
41.67
45.45
40.00
37.50
Nature of primary constriction
-
Nearly median
Nearly submedian
Remarks
Chromosome pair with secondary constriction.
-
-
- - - -
Fig. 14c Fig. 15c Fig. 16c
Fig. 18c
Fig. 14d Jt a8 88 88 81 w #I #I,
Fig. 16d 81 41 11 11 11 1) I! .!I 41 I J II 8 8 18, - Fig. 17d 41 81 11 I J I J JJ 81,
A C D
Fig. 18, @ 11 11 l t 11 - 11 jd dd If, 41 81 - 81 11, K c. D
Figs. 14c, 14d - Karyotype and idiogram of Acrocephalus capitatus Benth. (2n=18) x 2400 approx.; 15c, 15d - Karyotype and idiogram of Orthosiphon glabratus Benth. ( 2 ~ 2 6 ) x 2400 approx., 16c, 16d - Karyotype and idiogram of 2. glabratus var. parviflorus Benth. (2n=26) x 2400 approx., 17c, 17d - karyotype and idiograrn of 0.grandiflorus Bold cv.lilac (2n=28) x 2400 approx.; 18c, 18d - Karyotype and - - idiogram of - 0. srandiflorus Bold cv. white (2n=28) x 2400 approx.
Orthosiphon pallidus Royle (2n=4x=28=A4C8D16)
Chromosome count on pollen mother cell
Normal somatic chromosome number
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (Z)
n = 14 (Fig. 19j)
2n = 28 (Figs. 19b, 19c)
2
2.6 p to 1.2 p
1.73 P.I
48.4 P.I
12.1 P.I
36.84
25.33
39.68
Table 19 -
Detailed Karyotype analysis (Fig. 19d)
Chromo- some type
A
A
C
D
D
No. of pairs
1
1
4
4
4
I I I I I
Total length in P.I
2.6
2.4
2.0
1.6
1.2
Short arm length in pl
0.6 0.8
0.6 0.8
0.8
0.6
0.4
F%
23.08 30.77
25.00 33.33
40.00
37.50
33.33
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Remarks
Chromosome pair with secondary constriction.
-
- -
Orthosiphon thymiflorus Roth. (2n=4x=24=A4C20)
Chromosome count on pollen mother cell
Normal somatic chromosome number
Chromosome pairs with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
n = 12 (Fig. 20j)
2n = 24 (Figs. 20b, 20c)
2
3.6~ to 2.0~
2.53~
60.8~
15.2~
28.57
15.59
45.07
Table 20 --
Detailed taryotype analysis (Fig. 20d)
Remarks
Chromosome pair with secondary constriction
-
-
-
-
-
-
Chromo- some type
A
A
C
C
C
C
C
C
F%
22.22 27.78
21.43 28.57
42.86
38.46
33.33
41.67
36.36
40.00
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
Nearly submedian
Nearly median
No. of pairs
1
1
1
2
3
1
2
1
Total length in p
3.6
2.3
2.8
2.6
2.4
2.4
2.2
2.0
Short arm length in p
0.8 1 .'O
0.6 0.8
1.2
1.0
0.8
1.0
0.8
0.8
Plectranthus numrmularius Briq. (2n=4x=28=A4D24)
Chromosome count on pollen mother cell
Normal somatic chromosome number
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
n = 14 (Fig. 21j)
2x1 = 28 (Figs. 21b, 21~)
2
1.8 p to 1.0 ,u
1.34 p
37.6 ,u
9.4 p
28.57
19.89
40.57
Table 3
Detailed Karyotype analysis (Pig. 21d)
Remarks
Chromosome pair with secondary constriction.
-
-
-
-
Chromo- some type
A
D
D
D
D
Total length in p
1.8
1.6
1.4
1.2
1.0
No. of pairs
2
2
3
4
3
Short arm length in p
0.4 0.6
0.6
0.6
0.4
0.4
F%
22.22 33.33
37.50
42.86
33.33
40.00
Nature of primary constriction
-
Nearly submedian
Nearly median
Nearly submedian
Nearly median
--
F i g . 17b F i g . 1 7 j - - -
Fig. 18b
-.
Fig. 18j Fig. 19b Fig. 19j
- I F i g . 20b
- -- - - Fig. 2 0 j Fig. 21b
Scale : - g*
Figs. 17b, 1 7 j - Orthosiphon grandiflorus Bold. cv. lilac: 17b - mitotic metaphase (2n=28), 17j - meiotic metaphase 1 (11~14); 18b, 18j - - 0. grandiflorus Bold. cv. white: 18b - mitotic rnetaphase (2n=28), 18j - -- diakinesis (n=14); 19b, 1 9 j - 0, pallidus Royle: 19b - mitotic metaphase (2n=28), 1 9 j - meiotic metaphaso-I (n=14); 20b, 2 0 j - 0. thymiflorus Roth.: 20b-, mitotic metaphase (2n=24), 20j - meiotic anaphase I (n=12); 21b - Plectranthus nummularius B r i q . - mitotic metaphase (2n=28).
Plectranthus wightii Benth.(2n=4~=24=A4C4DI6)
Chromosome count on pollen mother cell
Normal somatic chromosome number
Chromosome pairs with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
n = 12 (Fig. 22j)
2n = 24 (Figs. 22b, 22c)
2
2.2 p to 1.4 p
1.78 p
42.8 p
10.7 p
22.22
12.53
42.56
Table 22 -- Detailed karyotype analysis (Fig. 22d)
Remarks
Chromosome pair with secondary constriction
-
-
-
-
I
Chromo- some type
A
A
C
D
D
D
Short arm length in p
0.4 0.8
0.4 0.6
0.8
0.8
0.6
0.6
I
No. of pairs
1
1
2
3
4
1
I
F%
18.18 36.36
20.00 30.00
40.00
44.44
37.50
42.86
I
Total length in p
2.2
2.0
2.0
1.8
1.6
1.4
I
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
I
Coleus arematicus Benth. (2n=4x=32=A4C22D6)
Normal somatic chromosome number
Somatic variation numbers
Chromosome pair with secondary constriction
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%I
: 2n = 32 (Figs. 23b, 23c)
: 2n = 16, 24, 30, 34 and 48 (Figs. 23e, 23f ,23g, 23h,23i)
Table 23 -- Detailed karyotype analysis (Fig. 233)
Remarks
Chromosome pair with secondary constriction
- -
- -
-
Nature of primary constriction
-
-
Nearly median
Nearly submedian
F%
20.00 33.33
23.08 30.77
42.86
41.67
45.45
44.44
37.50
Chromo- some type
A
A
C
C
C
D
D
Total length in p
3.0
2.6
2.8
2.4
2.2
1.8
1.6
No. of pairs
1
1
2
3
6
1
2
Short arm length in p
0.6 1.0
0.6 0.8
1.2
1.0
1.0
0.8
0.6
F i g s . 19c
Fig . 21c
Fig . 20c
F ig . 22c
F ig . 22d 88 8Cf l 8 8 18 8 l 11 o 81 8 1 18, A C D
F i g s . 19c, 19d Karyotype and id iogram of Orthosiphon p a l l i d u s Royle (211~28) x 2400 approx. ; 20c, 20d - Karyotype and idiograrn of 0 . t h y o i f l o r u s Roth . ( 2 ~ 2 4 ) x 2400 approx . ; 21c, 21d - Karyotype and idiogram,f P l e c t r a n t h u s - nummularius Br iq . (2n=28) x 2400 approx . , 22c 22d - Karyotype and idiograrn of - P. w 9 h t i i Benth. (2n=24; x 2400 approx .
Coleus aromaticus Benth. var. variegata (2n=4x=32=A4C16D12)
Normal somatic chromosome number
Somatic variation numbers
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value ( % I
2n = 32 (Figs 24b, 24~)
2u = 16, 24, 30 and 44 (Figs.Z4e,24f,24g,24h)
Table 24 -- Detailed karyotype analysis (Pig.24d)
Remarks
Chromosome pair with secondary constriction
-
-
-
-
Chromo- some type
A
A
C
C
C
D
D / 3 1 . 6 0.6 I 3 7 . 5 0 1 ~ .. ~- -- __I_. __ I -
No.of pairs
-
1
1
1
3
4
3
Total length in p
2.8
2.4
2.4
2.2
2.0
1.8
Shortarm length in p
0.6 1.0
0.6 0.8
0.8
0.8
0.8
0.6
F%
21.43 35.71
25.00 33.33
33.33
36.36
40.00
33.33
Nature of primary constriction
-
-
Nearly submedian
Nearly median
Nearly submedian
Fig. 23c
Fig. 23h
Fig. 23e
Fig. 23i
Fig. 23f
Fig. 24c
Figs. 23c, 23d. 23e, 23f. 23h, 23i - Coleus aromaticus Benth.: 23c, 23d - Karyotype and idiogram (2n=32) x 2400 approx.;23e, 23f, 23h, 23i - somatic variants with 2n=16, 2n=24, 2 ~ 3 4 and 2n=48 respectively; 24c, 24d - karyotype and idiogram of - C. aromaticus Benth. var. variegata (2n=32) x 2400 approx.
Coleus bl-i Benth. var. verschaffeltii (2n=4x=48=A4C8D36)
Normal somatic chromosome number
Somatic variation numbers
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
2n = 48 (Figs. 25b, 2%)
2n=46 (Fig. 25e)
2
2.6 p to 1.2 p
1.76 p
84.4 I.I
21.1 p
36.84
17.65
41.44
Table 25 -- Detailed karyotype analysis (Fig. 2 M )
Remarks
Chromosome pair with secondary constriction
- -
-
-
-
-
-
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearlymedian
Nearly submedian
Chromo- some type
A
A
C
D
D
D
D
D
D
Short arm length in p
0.6 0.8
0.6 0.8
0.8
0.8
0.6
0.6
0.6
0.4
0'. 4
F%
23.08 30.77
25.00 33.33
40.00
44.44
33.33
37.50
42.86
28.57
33.33
No. of pairs
1
1
4
7
1
5
3
1
1
Total length in p
2.6
2.4
2.0
1.8
1.8
1.6
1.4
1.4
1.2
Fig. 21j Fig. 22b F i g . 22j
- - - -- Fig. 23b Fig. 23g Fig. 2 4 7 --
P
Fig. 24f --
F i g . 24h Fig. 25b
Fig. 21j - Plectranthus nummularius Briq. - meiotic metaphase I (abnormal with 13 bivalents and 2 univalents); 22b, 22j - P. wightii Benth.: 22b - mitotic metaphase (2n=24), 22 j - meiotic anaphase (n=i2); 23b, 23g - Coleus aromaticus Benth. : 23b -mitotic metaphase (2n=32), 23g - somatic variant (2n=30) ; 24b , 24f, 24h - C. aromaticus Benth. var. variegata : 24b - mitotic metaphase (2n=32),, 24r - somatic variant (2n~24); 24h - somatic variant (2n=44); 25b - - C. blumei Benth. var. verschaffeltii - mitotic metaphase (2nZ48).
Coleus forskoblii Briq. (2n=4x=28=A4C22D2)
Normal somatic chromosome number
Somatic variation numbers
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
2n = 28 (Figs. 26b, 26c)
2n = 26 and 30 (Figs. 26e, 26f)
2
3.4 p to 1.6 u
2.36 ,u
66 P
16.5 ,u
36
17.89
42.11
Table 26 --
Detailed Karyotype analysis (Pig. 26d)
Chromo- some type
A
A
C
C
C
C
No.of pairs
1
1
1
5
2
3
D I 1
/ 1.6 / 0.6 / 37.50 / Nearly submedian -
Remarks
Chromosome pair with secondary constriction
-
-
-
-
Total length in p
3.4
2.8
2.8
2.4
2.2
2.0
Shortarm length in u
0.6 1.0
0.6 0.8
1.0
1.0
1.0
0.8
F%
17.65 29.41
21.43 28.57
35.71
41.67
45.45
40.00
Nature of primary constriction
-
-
Nearly submedian
Nearly median
Fig. 24c
Fig. 25e
Fig. 24g
- 3 \I Fig. 26c
Fig. 25c
Fig. 26e
~ --
Figs. 24e,24g - Somatic variants of Coleus aromaticus Benth. var. variegata with 2n=16 and 2n=30 respectively; 25c,25d,25e - - C -- blumei Benth. var. verschaffeltii : 25c, 25d - Karyotype and idiogram ( 2 ~ 4 8 ) x 2400 approx., 25e - somatic variant (2~46); 26c, 26d, 26e - C. forskohlii Briq.: 26c, 26d - Karyotype and idiogram ( 2 ~ 2 8 ) x 2400 approx., 26e1 somatic variant (2n=26)
Coleus lacidatus Benth. (2n=4x=48=A4D44)
Normal somatic chromosome number
Somatic variation numbers
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length .
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
2n = 48 (Figs. 27b, 27~)
2n = 42 (Pig. 27e)
2
2.0 p to 1.2 p
1.49 p
Table 27 -- Detailed karyotype analysis (Fig. 27d)
Chromo- some type
A
A
D
D
D
D
No. of pairs
1
1
2
5
12
3 I
Total length in p
2.0
1.8
1.8
1.6
1.4
1.2 I
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
Nearly submedian I
Remarks
Chromosome pair with secondary constriction
-
-
-
- I
Short arm length in p
0.4 0.6
0.4 0.6
0.8
0.6
0.6
0.4 I
F%
20.00 30.00
22.22 33.33
44.44
37.50
42.86
33.33 I
5 5
Coleus parviflorus Benth. (Zn=6x=72=AbD66)
Normal somatic chromosome number
Somatic variation numbers
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
2n = 72 (Figs. 28b, 28c)
2n = 24,32,36,48 and 68 (Figs. 28e,28f ,28g,28h,281)
Table 28 --
Detailed karyotype analysis (Fig. 28d)
Chromo- some type
A
A
D
D
D
D
No.of pairs
2
1
1
6
22
4
Total length in p
2.0
1.8
1.8
1.6
1.4
1.2
Shortarm length in p
0.6 0.6
0.4 0.6
0.6
0.6
0.6
0.4
F%
30.00 30.00
22.22 33.33
33.33
37.50
42.86
33.33
Nature of primary constriction
-
-
Nearly submedian
Nearly median
Nearly submedian
Remarks
Chromosome pair with secondary constriction
-
-
- -
Fig. 26f Fig. 27c Fig. 27e
Fig. 28c Fig. 28e Fig. 28g
Fig. 26f - Somatic variant of Coleus forskohlii Briq. with 2n=30; 27c,27d,27e - C.laciniatus Benth.: 27c, 27d - Karyotype and idiogram (2n=48) x 2400 approx., 27e - - somatic variant with 2n=42; 28c, 28d, 28e, 28g - Coleus parviflorus Benth.: 28c 2nd - Karyotype and idiogran (2n=72) x 2400 approx. 2 8 e 288 - somatic variants with 2n=24 and 2n=36 respectively
Coleus rehneltianus Berger (Zn=4~=48=A4CZD42)
Normal somatic chromosome number
Somatic variation numbers
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
2n = 48 (Figs. 29b, 29c)
2n=50 (Fig. 29e)
2
2.2 p to 1.2 p
1.6 p
76.8 p
19.2 p
29.41
15.73
40.33
Table 29 --
Detailed karyotype analysis (Pig. 29d)
Chromo- some type
A
A
C
D
D
D
D
D
No. of pairs
1
1
1
3
2
6
8
2
Total length in p
2.2
2.0
2.0
1.8
1.8
1.6
1.4
1.2
Short arm length in p
0 .'4 0.8
0.4 0.6
- 0.8
0.6
0.8
0.6
0.6
0.4
F%
18.18 36.36
20.00 30.00
40.00
33.33
44.44
37.50
42.86
33.33
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nerarly median
Nearly submedian
Nearly median
Nearly submedian
Remarks
Chromosome pair with secondary constriction
-
-
-
-
-
-
Coleus zeylanicus (Benth.) Cramer (2n=4x=28=A4C8D16)
Normal somatic chromosome number
Somatic variation numbers
Chromosome pair with secondary constriction
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
2n = 28 (Figs. 30b, 30c)
2n = 14, 30, 32 and 56 (Figs. 30e, 30f, 30g, 30h)
Table 30 --
Detailed karyotype analysis (Fig. 30d)
Chromo- some type
A
A
C
C
D
D
D
No.of pairs
1
1
2
2
4
2
2
I
Total length in p
2.8
2.2
2.2
2.0
1.8
1.6
1.4
I
Nature of primary constriction
-
-
Nearly submedian
Nearly median
Nearly submedian
Nearly median
I
Remarks
Chromosome pair [with secondary constriction
-
-
-
-
Shortarm length in p
0.6 0.8
0.4 0.8
0.8
0.8
0.8
0.6
0.6
I
F%
21.43 28.57
18.18 36.36
36.36
40.00
44.44
37.50
42.86
I
Fig. 28i @ Fig. 29c Fig. 29e
Fig. 30c Fig. 30e Fig. 30f
Fig 281 Sonatic variant of - Coleus E--. arviflorus Benth. (2~68); 29c,29d,29e C rehneltianus Berzer 29c,29d - Karyotype and idiogram (2n=48) x 2400 approx., 29e - somatic variant (2n-50); 30c,30d:30e,30f - C. zeylanicus .-
(Benth., Cramer 30c, 30d - Karyotype and idiogram (2n-28, x 2400 appro~. , 30e 30f somatic variants with 2n=30 and 2n=32 respectively.
dnisochilus carnosus ( L . f . ) Wall. (2n=Zx=34=A4C14D16)
Normal somatic chromosome number
Chromosome pair with secondary constriction
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
: 2n = 34 (Figs. 31b, 31c)
: 2
: 3.4,u to 1.2p
: 2.1@
: 73.61~
: 36.41
: 47.83
: 30.00
: 42.24
Table 31 -- Detailed karyotype analysis (Fig. 31d)
Chromo- some type
A
A
C
C
C
D
D
D
D
F%
17.65 29.41
18.75 31.25
42.86
41.67
40.00
44.44
37.50
42.86
33.33
No. of pairs
1
1
3
3
1
3
2
2
1
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
Nearly submedian
Total length in p
3.4
3.2
2.8
2.4
2.0
1.8
1.6
1.4
1.2
Remarks
Chromosome pair with secondary constriction
-
-
- -
-
- -
Short arm length in j~
0.6 1.0
0.6 1.0
1.2
1.0
0.8
0.8
0.6
0.6
0.4
A n i s ~ u s eriocepbalus Benth. (2n=2x=32=A4C12D16)
Normal somatic chromosome number
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (Dl)
Variation coefficient (VC)
TF value (%)
2n = 32 (Figs. 32b, 32c)
2
3 . 2 p to 1.2 p
1 .91 p
61.2 u
30.6 p
45.45
28.91
41.45
Table 32 --
Detailed karyotype analysis (Fig. 32d)
Chromo- some type
A
A
C
C
C
D
D
D
No. of pairs
1
1
2
1
3
3
4
1
Total length in p
3.2
2.8
2.4
2 .2
2 .0
1.6
1.4
1.2
Short arm length in &I
0.8 , 1 . 0
0 . 6 0 .8
1 . 0
0 .8
0 .8
0 . 6
0.6
0 .4
F%
25.00 31.25
21.43 28.57
41.67
36.36
40.00
37.50
42.86
33.33
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
Nearly submedian
Nearly median
Nearly submedian
Remarks
Chromosome pair with secondary constriction
- -
-
- -
--- Fig. 26b
. * ', 'v'
Fig. 27b
- ~ - -~
Fig. 28f Fig. 28h Fig. 29b
-- -. --
~ .-
Fig. 30b
T( f 9 / it' ,R/ 3 4
Fig. 31b -
Fig. 32b
Scale: - SP
Fig. 26b - Coleus forskohlii Briq. - mitotic metaphasc (211'281, 27b-C. lachiatus Benth. - mitotic metaphase (2n=48); 28b, 28f ; X h - C. parviflor& Benth.: 28b - mitotic metaphase (2nZ72), 28f - somatic variant Dn=32), 28h - somatic variant (2n=48); 29b - C. rehnelcianus Serger - mitotic metaphase (2n=48); 30b - C. zeylanicus (~eni-h.) Cramer - sitotic metaphase (2n=28); 31b - Anisochilus carnosus (L . f . ) Wall. - mitotic mctaphase (2n=34); 32b - - A. - eriocephalus Benth. - mitotic metaphase (?n=32).
Eyptis capitata Jacq. (2n=4x=32=A4D28)
Normal somatic chromosome number : 2n = 32 (Figs. 33b, 33c)
Chromosome pair with secondary constriction : 2
Range of chromosome length : 1.81-1 to 1.0~
Average chromosome length : 1.35~
Total chromatin length : 43.q~
Chromatin length of basic complement : 10.8~
Disparity index (DI) : 28.57
Variation coefficient (VC) : 19.25
TF value (%) : 39.68
Table 33 --
Detailed taryotype analysis (Pig. 33d)
Chromo- some type
A
D
D
D
D
D
No. of pairs
2
1
1
3
7
2
Total length in p
1.8
1.8
1.6
1.4
1.2
1.0
Short arm length in p
0.4 0.6
0.8
0.6
0.6
0.4
0.4
F%
22.22 33.33
44.44
37.50
42.86
33.33
40.00
Nature of primary constriction
-
Nearly median
Nearly submedian
Nearly median
Nearly submedian
Nearly median
Remarks
Chromosome pair with secondary constriction
-
-
- -
-
Fig. 30g
Fig. 32c
& 4 - \ #
Fig. 30h
Fig. 31c
Fig. 33c
33d &jp,Jt I t I# rr #I rt r ~ ~ t #I JI 18 J# B, ,b --
Figs 30g, 30h - - Coleus . zeylanicus (Benth.) Cramer: 30g - somatic variant (2nz32), 30h - somatic variant (2n=56); 31c, 31d - karyotype and idiogram of Anisochilus carnosus ( L . f . ) Wall. ( 2 ~ 3 4 ) x 2400 approx.;32c, 32d - Karyotype and idiogram of A. eriocephalus Benth. (2n=32) x 2400 approx.; 33c, 33d - Hyptis capitata Jacq. - (2n=32) x 2400 approx.
Normal somatic chromosome number
Chromosome pair with secondary constriction , :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
2n = 28 (Figs. 34b, 34c)
2
1.6 p to 1 . 0 p
1 . 3 p
36 .4 p
9.1 p
23.08
12.67
39.04
Table 31r -- Detailed karyotype analysis (Fig. 34d)
Chromo- some type
A
A
D
D
D
D
I I I I I
No. of pairs
1
1
1
3
7
1
Total length in p
1.6
1.4
1.6
1.4
1.2
1.0
Short arm 'length in p
0.4 0 .4
0 .4 0.4
0.6
0.6
0 .4
0 .4
Nature of primary constriction
-
-
Nearly submedian
Nearly median
Nearly submedian
Nearly median
F%
25.00 25.00
28.57 28.57
37.50
42.86
33.33
40.00
Remarks
Chromosome pair with secondary constriction
-
- -
-
Pogostemon benghalensis (Burm.f.) Kuntze (2n=4x=64=A4D60)
Normal somatic chromosome number
Somatic variation numbers
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value ( X )
2n = 64 (Figs. 35b, 35c)
32 and 66 (Figs. 35e, 35f)
2
1.8 p to 1.0 p
1.24 cl
79.6 pl
19.9 p
28.57
14.94
37.77
Table 35 --
Detailed karyotype analysis (Pig. 3%)
Chromo- some type
A
A
D
D
D
D
I l l I I
No. of pairs
1
1
2
5
17
6
Total length in p
1.8
1.4
1.6
1.4
1.2
1.0
Short arm length in w
0.4 0.6
0.4 0.4
0.6
0.6
0.4
0.4
Nature of primary constriction
-
-
Nearly submedian
Nearly median
Nearly submedian
Nearly median
F%
22.22 33.33
28.57 28.57
37.50
42.86
33.33
40.00
Remarks
Chromosome pair with secondary constriction
-
-
-
-
Pogostemon heyneanus Benth. (2n=4x=64=A4D60)
Normal somatic chromosome number
Somatic variation numbers
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
2n = 64 (Figs 36b, 36c)
2n=68 (Fig. 36e)
2
Table 36 -- Detailed karyotype analysis (Fig. 36d)
Chromo- some type
A
A
D
D
D
D
I l l I I I
No.of pairs
1
1
7
7
12
4
Total length in p
1.8
1.6
1.6
1.4
1.2
1.0
Shortarm length in p
0.4 0.6
0.4 0.4
0.6
0.6
0.4
0.4
F%
22.22 33.33
25.00 25.00
37.50
42.86
33.33
40.00
Nature of primary constriction
-
-
-
Nearly submedian
Nearly median
Nearly submedian
Nearly median
Remarks
Chromosome pair with secondary constriction
- -
- -
Fig. 34c
B -
Fig. 35f
Fig. 35c
Fig. 35e
Fig. 36e
Fig. 34d88 48 81 11 11 11 I1 I 1 81 I t I 1 18 II I! -\
Fig.34c, 34d - Karyotype and idiogram of Hyptis suaveolens ( L . ) Poit. (2n=28) x 2400 approx.; 35c, 35d, 35e, 35f - Pogostemon benghalensis (Burm.f.) Kuntze : 35c, 35d - Karyotype and idiogram (2n=64) x 2400 approx., 35e - somatic variant (2n=32), 35f - somatic variant (2n=66j; 36c, 36d, 36e - P. heyneanus Benth.: 36c, 36d - Karyotype and idiogram i2n=64) x 2400 apcrox., 36e - somatic varlant (211~68)
Pogosteron purpurascens Dalz . (2n=Zx=32=A4CZD26)
Normal somatic chromosome number : 2n = 32 (Figs. 37b, 37c)
Somatic variation numbers : 2n = 30 (Fig. 37e)
Chromosome pair with secondary constriction : 2
Range of chromosome length : 2.2 p to 1.2 p
Average chromosome length : 1.58 p
Total chromatin length : 50.4 p
Chromatin length of basic complement : 25.2 p
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
Table 37 --
Detailed karyotype analysis (~ig. 37d)
Chromo- some type
A
A
C
D
D
Total length in p
2.2
2.0
2.0
1.6
1.4
No. of pairs
1
1
1
5
7
1
I I I
Short arm length in p
0.4 0.6
0.4 0.6
0.8
0.6
0.6
33.33
O.* I I I
F%
18.18 27.27
20.00 30.00
40.00
37.50
42.86
Nearly submedian -
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
Remarks
Chromosome pair with secondary constriction
-
-
-
Pogostenon vestitus Benth . (2"=2~=32=A4D28)
Normal somatic chromosome number
Somatic variation numbers
Chromosome pair with secondary constriction
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
2n = 32 (Figs. 38b, 38c)
34 (Fig. 38e)
2
2.0 p to 1.0 p
1.39 p
44.4 p
22.2 p
33.33
20.01
40.53
Table 38 --
Detailed karyotype analysis (Fig. 38d)
Remarks
Chromosome pair with secondary constriction.
-
- -
D I 2 ( 1.0 / 0.4, /4O.OO/Nearlymedlan 1 -
Short arm length in p
0.4 0.6
0.4 0.6
0.8
0.6
0.6
0.4
Total length in p
2.0
1.8
1.8
1.6
1.4
1.2
Chromo- some type
A
A
D
D
D
D
No. of pairs
1
1
1
1
5
5
F%
20.00 30.00
2 2 . 2 2 33.33
44.44
37.50
42.86
33.33
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearlymedian
Nearly submedian
6 6
Zusteralis quadrifolia (Benth3 Panigrahi (2n=2x=30=A4D26)
Normal somatic chromosome number
Chromosome pair with secondary constriction
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value ( % I
2n = 30 (Figs. 39b, 3%)
2
1.6 ,u to 1.0 p
1.16 p
34.8 p
17.4 p
23.8
16.89
38.67
Table 39 - -
Detailed Karyotype analysis (Fig. 39d)
Chromo- some type
A
D
D
No.of pairs
2
6
7
Total length in p
1.6
1.2
1.0
Shortarm length in p
0.4 0.4
0.4
0.4
F%
25.00 25.00
33.33
40.00
Nature of primary constriction
-
Nearly submedian
Nearly median
Remarks
Chromosome pair with secondary constriction.
-
-
Fig. 38c
Fig. 37c Fig. 37e
Fig. 38e Fig. 39c
Fig. 39d $8 ,tt 11 81 18 81 18 11 # t 11 11 11 11 11, /
- - A D
Figs. 37c 37d, 37e Pogostemon Krpurnscens Dalz.: 37c, 37d - karyotype and -- - -- idiogram (2n=32) x 2400 approx., 37e - somatic variant (2~30); 38c, 38d. 3& - P. vestitus Benth.: 38c, 38d - Karyotype and idiogram (2n=32) x 2400 - approx., 38e - somatic variant (2n=34); 39c, 39d - Karyotype and idiogram of Eusteralis quadrifolia (Benth.) Panigrahi !2n=30) x 2400 approx.
Normal somatic chromosome number
Somatic variation numbers
Chromosome pair with secondary constriction
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
2n = 96 (Figs. 40b, 40c)
2n=72 (Fig. 40e)
2
1.8 p to 1.0 p
1.17 p
112.4 p
14.05 p
28.57
17.79
37.98
Table 40 -- Detailed karyotype analysis (Fig. 40d)
Total lengtt in P
Chromo- some type
No. of pairs
Short arm length in ~ . r
Chromosome pair with secondary constriction
Nature of primary constriction
F% Remarks
Nearly submedian
Nearly median
Nearly submedian
-
- -
1 Nearly median - -
- -- -- -
Fig. 33b
-
Fig. 36b
-
Fig. 34b
-- --
Fig. 37b
Fig. 35b
Fig. 38b
Fig. 39b Fig. 40b Fig. 40e
Scale : - G P
Fig. 33b - Hyptis capitata Jacq. - mitotic metaphase (2n=32); 34b - H. suaveolens ( L . ) Poit. - mitotic metaphase (2n = 28); 35b - ~ogostemon benghalensis (Burm.f.1 Kuntze - mitotic metaphase (211~64); 36b - P. heyneanus Benth. - mitotic metaphase (2n=64); 37b - - P. purpurascens ~alz: - mitotic netaphase (2n=32); 38b - P. vestitus Benth. - mitotic metaphase (2~32); 39b - Eusteralis quadrifolia T~enth. ) Panigrahi - mitotic metaphase (2ns30) ; 40b, 40e - Mentha arvensis L.: 40b - mitotic metaphase (2n=96); 40e - somatic variant (2n=72);
Mentha rotuudifolia (L.) Huds.(211=2~=24=A4D20)
Normal somatic chromosome number
Somatic variation numbers
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
2n = 24 (Figs. 41b, 41c)
2n=36 and 48 (Figs. 41e. 41f)
2
Table 41 -- Detailed karyotype analysis (Pig. 41d)
Chromo- some type
A
A
D
D
No. of pairs
1
1
2
8
Total length in p
1.6
1.4
1.2
1.0
Short arm length in p
0.4 0.4
0.4 0.4
0.4
0.4
F%
25.00 25.00
28.57 28.57
33 .33
40.00
Nature of primary constriction
-
-
Nearly submedian
Nearly median
Remarks
Chromosome pair with secondary constriction
- -
llentha spicata(L.) Huds. (2n=4x=48=A4D44)
Normal somatic chromosome number : 2n = 48 (Figs. 42b, 42c)
Somatic variation numbers : 2n=24, 64 and 72 (Figs. 42e, 42f, 42g)
Chromosome pair with secondary constriction : 2
Range of chromosome length : 1.8 p to 1.0 p
Average chromosome length : 1.08 p
Total chromatin length : 51.6 p
Chromatin length of basic complement : 12.9 p
Disparity index (DI) : 28.57
Variation coefficient (VC) : 16.75
TF value (%) : 40.53
Table 42 --
Detailed karyotype analysis (Pig. 42d)
Chromo- some type
A
A
D
D
No. of pairs
1
1
3
19
Total length in p
1.8
1.4
1.2
1.0
Short arm length in p
0.4 0.6
0.4 0.4
0.4
0.4
F%
22.22 33.33
28.57 28.57
33.33
40.00
Nature of primary constriction
-
-
Nearly submedian
Nearly median
Remarks
Chromosome pair with secondary constriction
-
-
Fig. 40c Fig. 41c
Fig. 41f Fig. 42c
Fig. 41e
Fig. 42e
Fig. 40c,40d - Karyotype and idiogram of Mentha arvensis L.(2n=96) x 2400 approx.; 41c 41d,41e 41f - - M. rotundifolia (L.) Huds.: 41c,41d - Karyotype and idiogram (2n=24> x 2400 approx , 41e, 41f - somatic variants with 2n=36 and 2n=48 respectively.; 42c,42d,42e - M. scats (L. ) Huds.: 42c, 42d - Karyotype and - -. idiogran ( 2 ~ 4 8 ) x 2400 approx., 42e somatic variant with 2n=24
S a l a coccinea Juss. (2n=Zx=22=A2C8D12)
Chromosome count on pollen mother cell
Normal somatic chromosome number
Somatic variation numbers
Chromosome pair with secondary constriction :
Range of Chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
n = 11 (Fig. 433)
2n = 22 (Figs. 43b, 43c)
2n = 20 and 24 (Figs. 43e, 43f)
1
2.6 p to 1.6 p
1.95 p
42.8 p
21.4 p
23.81
18.08
37.92
Table 43 --
Detailed Karyotype analysis (Fig. 43d)
Nature of primary constriction
-
Nearly submedian
Nearly median
Nearly submedian
Chromo- some type
A
C
C
D
D
Remarks
Chromosome pair with secondary constriction.
-
- -
-
Total length in fl
2.6
2.4
2.0
1.8
1.6
No. of pairs
1
2
2
2
4
Short arm length in p
0.6 0.8
0.8
0.8
0.6
0.6
F%
23.07 30.77
33.33
40.00
33.33
37.50
Salvia leucantha Cav. (2n=2x=22=A2C18D2)
Chromosome count on pollen mother cell
Normal somatic chromosome number
Somatic variation numbers
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length .
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
n = 11 (Fig. 44j)
2n = 22 (Figs. 44b, 44~)
2n = 23 and 24 (Figs. 44e, 44f)
1
2.8 p to 1.6 p
2.47 p
54.4 !J
27.2 ,u
27.27
14.35
40.95
Table 3 - Detailed Karyotype analysis (Pig. 44d)
-- - . . ,
Fig. 41b Fig. 42b Fig. 42g
- - -- --
Fig. 43b Fig. 43e Fig. 43j
Fig. 44b Fig. 44e Fig. 44 j
Scale: U BP
Fig. 41b - Mentha rotundifolia ( L . ) Huds, - mitotic metaphase, (2n=24); 42b, 42g - M. spicata (L.) Huds.: 42b - mitotic metaphase (2n=48), 42g - somatic variant (2n=72); 43b, 43e, 43j - Salvia coccinea Juss.: 43b - mitotic metaphase (2n=22), 43e - somatic variant (2n=20), 43 j - diakinesis (n=ll); 44b, 44e, 44j - S. leucantha Cav.: 44b - mitotic metaphase (2n=22), 44e - somatic varia.nt (%=23), 441 - meiotic anaphase I (n=ll)
Salvia plebeia R.Br. (2n=2~=16+0-IB=A2C4DlO)
Chromosome count on pollen mother cell : n = 8+0-1B (Fig. 453)
Normal somatic chromosome number : 2n = 16+0-1B (Figs. 45b, 45c)
Somatic variation numbers
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
2n=14 & 18 (Figs. 45e, 45f)
1
2.6 ,u to 1.4 p
1.83 p
29.2 p
Table 45 -- Detailed karyotype analysis (Fig. 45d)
Chrono- some type
A
C
D
D
D
No. of pairs
1
2
1
3
1
Total length in p
2.6
2.0
1.8
1.6
1.4
Short arm length in p
0.6 0.8
0.8
0.6
0.6
0.6
F%
23.08 30.77
40.00
33.33
37.50
42.86
Nature of primary constriction
-
Nearly median
Nearly submedian
Nearly median
Remarks
Chromosome pair with secondary constriction
-
-
-
-
Fig. 42f
Fig. 44c
Fig. 43c Fit.43f
Fig. 44f
Fig. 43d & II I I 11. .II ir JI 18 rt rr A C D
Fig. 44d
Fig. 45d
Fig. 45c
Fig. 42f - -- Mentha spicata ( L . ) Huds.-somatic variant (2n=64); 43c, 43d, 43f - Salvia coccinea Juss.: 43c, 43d - karyotype and idiogram i2n-22) x 2400 appxox., 43f - somatic variant (2n=24); 44c 44d, 44f - 5.leucantha Cav.: 44c, 44d - karyotype and idiogram (2n=22) x 2400 approx.; 44f - somatic variant (2n=24); 45c, 45d - karyotype and idiogram of plebeia R.Br. [2n=16+0-1 B :" B Chromosome)] x 2400 approx.
Salvia splendens ker - Gawl. (2n=4x=44=A4C2D38)
Chromosome count on pollen mother cell
Normal somatic chromosome number
Somatic variation numbers
Chromosome pairs with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
n = 22 (Fig. 46j)
2n = 44 (Figs. 46b, 46c)
2n=22 and 48 (Figs. 46e, 46f)
2
2.2 p to 1.2 p
1.63 p
71.6 p
17.9 p
29.41
17.07
41.19
Table 46 --
Detailed karyotype analysis (Pig- 46d)
Chromo- some type
A
A
C
D
D
D
D 1 3 1 1.2 / 0.4 I33.33/Nearlysubmedian I -
No. of pairs
1
1
1
5
7
4
Total length in p
2.2
2.0
2.2
1.8
1.6
1.4
Short arm length in p
0.6 0.6
0.4 0.6
1.0
0.8
0.6
0.6
Remarks
Chromosome pair with secondary constriction
-
-
-
-
F%
27.27 27.27
20.00 30.00
45.45
44.44
37.50
42.86
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
Anisomeles indica (LJ Kuntze (2n=Zx=34=A4D30)
Normal somatic chromosome number : 2n = 34 (Figs. 47b, 47c)
Chromosome pair with secondary constriction : 2
Range of chromosome length : 1.6 p to 1.0 p
Average chromosome length : 1.19 p
Total chromatin length : 40.4 p
Chromatin length of basic complement : 20.2 p
Disparity index (DI) : 23.08
Variation coefficient (VC) : 14.66
TF value (%) : 39.19
Table 47 -- Detailed Karyotype analysis (Pig. 47d)
Chromo- some type
No. of pairs
1
1
2
7
6
Total length in p
1.6
1.4
1.4
1.2
1.0
Short arm length in p
0 .'4 0.4
0.4 0.4
0.6
0.4
0.4
.
Remarks
Chromosome pair with secondary constriction
-
-
-
F%
25.00 25.00
28.57 28.57
42.86
33.33
40.00
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
Fig. 45e
Fig. 46e
Fig. 45f
Fig. 46f
Fig. 46c
Fig. 47c
Fig. 45e, 45f - Salvia plebeia R. Br.: 45e - somatic variant (2n=14), 45f - somatic variant (2n=18); 46c, 46d, 46e, 46f - S. splendens Ker - Gawl.: 46c, 46d - karyotype and idiogram (2n=44) x 2400 approx., 46e - somatic variant (2n~7.3; 46E - somatic variant (2~1~48); 4732, 47d - Anisomeles indica (L.) Kuntze.-karyotype and idiogram (in-34) x 2400 approx.
Normal somatic chromosome number
Chromosome pair with secondary constriction
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value ( X )
2n = 32 (Figs. 48b, 48c)
Table 48 --
Detailed karyotype analysis (Pig. 48d)
Remarks
Chromosome pair with secondary constriction
-
-
-
-
I I I I I I
F%
20.00 30.00
22.22 33 .33
44.44
42.86
33 .33
40.00
Short arm length in p
0 .4 0.6
0.4 0 .6
0.8
0.6
0 .4
0 .4
Chromo- some type
A
A
D
D
D
D
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
No. of pairs
1
1
2
4
4
4
Total length in p
2.0
1.8
1.8
1.4
1.2
1.0
Fig. 45b
- - - -
Fig . 463
- - - -
Fig. 45 j Fig. 46b
. - -
Fig. 47b F i g . 48b
- I--- - -- -- - -
Fig. 49b Fig. 50b Fig. 51b
F i g . 45b , 4 5 j - Salvia p l e b e i a R .Br . : 45b-mitotic metaphase [2n=16+0-1~ (* B chromosome) 1, 45j - meiotic anaphase I [n=8+0-16 (* B chromosome) 1; 46b, 4 6 j - S . s p l e n d e n s Ker. Gawl.: 46b - mitotic metaphase ( 2 ~ 4 4 1 , 46j - meiotic - metaphase (abnormal airi in^ of bivalents): 47b - Anisomeles indica (L.) Kuntze - - <
- . - mitotic metaphase (211~34); 48b. - A. malabarica R . B r . - mitotic metaphase (2n=32) ; 49b - Leucas aspera spreng. - mitotic metaphase ( 2 ~ 2 2 ) ; 50b - 4. -
cephalotes Spreng. - mitotic metaphase (2n = 2 2 ) ; 51b - - L. linifolia Spreng. - mitotic rnetaphase (2n=22)
Leucas aspera Spreng. (2n=Zx=22=A4D18)
Normal somatic chromosome number
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
2n = 22 (Figs. 49b, 49c)
2
2.4 p to 1.0 p
1.44 p
31.6 p
15.8 p
41.18
26.36
37.48
Table 49 --
Detailed Karyotype analysis (Fig. 4%)
Chromo- some type
A
A
D
D
D
D
I I I I I I
No. of pairs
1
1
1
4
2
2
Total length in p
2.4
1.8
1.6
1.4
1.2
1.0
Short arm lengfh in p
0.6 0.8
0.4 0.6
0.6
0.4
0.4
0.4
F%
25.00 33.33
22.22 33.33
37.50
28.57
33.33
40.00
Nature of primary constriction
-
-
Nearly submedian
Nearly median
Remarks
Chromosome pair with secondary constriction.
-
-
-
-
Le- cephalotes Spreng. (2n=Zx=22=A4D18)
Normal somatic chromosome number
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
2n = 22 (Figs. 50b, 50c)
2
2.4 p to 1.0 p
1.47 p
32.4 p
16.2 p
41.18
26.76
38.53
Table 50
Detailed Karyotype analysis (Pig. 50d)
I I I I I I
Nature of primary constriction
-
-
Nearly submedian
Nearly median
Chromo- some type
A
A
D
D
D
D
Remarks
Chromosome pair with secondary constriction.
-
- -
-
Short arm length in p
0.6 0.8
0.6 0.6
0.6
0.4
0.4
0.4
No. of pairs
1
1
2
2
4
1
F%
25.00 33.33
30.00 30.00
37.50
28.57
33.33
40.00
Total length in p
2.4
2.0
1.6
1.4
1.2
1 .O
Fig. 48d
Fig. 49d
Fig. 50d
Fig. 51d
Fig. 50c
Fig. 49c
Fig. 51c
Fig. 48c, 48d - Karyotype and idioeram of Anisomeles malabarica R.Br. ( 2 ~ 3 2 ) x 2400 approx.; 49c, 49d - Karyotype and idiogram of Leucas aspera Spreng. (211-22) x 2400 approx.; 50c, 50d - Karyotype and i d i o x o f L.cephalotes Spreng. (2n=22) x 2400 approx.; 51c, 51d - Karyotype and idiogram of L.linifolia Spreng. (Zn=22) x 2400 approx. .-
78
Leucas linifolia Spreng. (2n=2x=22=A4D18)
Normal somatic chromosome number
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
2n = 22 (Figs. 51b, 51c)
2
1.6 ,u to 1.0 ,u
1.27 ,u
28.0 p
14.0 ,u
23.08
16.82
37.95
Table 51 --
Detailed Karyotype analysis (Pig. 51d)
Chromo- some type
A
D
D
D
No. of pairs
2
1
6
2
Total length in ,u
1.6
1.6
1.2
1.0
Short arm length in ,u
0.4 0.4
0.6
0.4
0.4
F%
25.00 25.00
37.50
33.33
40.00
Nature of primary constriction
-
Nearly submedian
Nearly median
Remarks
Chromosome pair with secondary constriction.
-
-
-
Leucas stricta Benth. (2n=Zx=22=A4D18)
Normal somatic chromosome number : 2n = 22 (Figs. 52b, 52c)
Chromosome pair with secondary constriction : 2
Range of chromosome length : 2.0 p to 1 . 0 ~
Average chromosome length : 1.35 p
Total chromatin length : 29.6 p
Chromatin length of basic complement : 14.8 p
Disparity index (DI) : 33.33
Variation coefficient (VC) : 21.04
TF value (%) : 37.09
Table 52 --
Detailed Karyotype analysis (Pig. 52d)
Chromo- some type
A
A
D
D
D
D
No.of pairs
1
1
1
2
4
2
I I I I
Total length in p
2.0
1.6
1.6
1.4
1.2
1.0
Shortarm length in p
0.4 0.6
0.4 0.4
0.6
0.4
0.4
0.4
F%
20.00 30.00
25.00 25.00
37.50
28.57
33.33
40.00
Nature of primary constriction
-
-
Nearly submedian
Nearly median
Remarks
Chromosome pair with secondary constriction.
-
-
-
-
Leucas vestita Benth. (2n=2~=22=A2ClODlO)
Normal somatic chromosome number
Chromosome pair with secondary constriction
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (XI
2n = 22 (Figs. 53b, 53c)
1
2.4 p to 1.4 p
1.95 p
42.8 p
21.4 p
26.32
14.56
43.64
Table 53 -- Detailed karyotype analysis (Fig. 5M)
Chromo- some type
A
C
C
D
D
D
NO. of pairs
1
3
2
3
1
1
Total length in p
2.4
2.2
2.0
1.8
1.6
1.4
Short arm length in p
0.4 0.8
1.0
0.8
0.8
0.6
0.6
F%
16.67 33.33
45.45
40.00
44.44
37.50
42.86
Nature of primary constriction
-
Nearly median
Nearly submedian
Nearly median
Remarks
Chromosome pair with secondary constriction
-
-
-
-
-
Leonotis nepetifolia (L.) Ait.f. (2n=4x=28=A2C24D2)
Normal somatic chromosome number
Chromosome pair with secondary constriction
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
: 2n = 28 (Figs. 54b, 54c)
: 1
Table 54 --
Detailed Karyotype analysis (Fig. 54d)
Chromo- some type
A
C
C
C
C
C
Short arm length in IJ
0.8 1.2
1.2
1.2
1.0
1.0
0.8
No. of pairs
1
1
1
4
4
2
D I 1 1.6 I 0.6 I 37.50 Nearly submedian
Total length in IJ
3.6
3.2
2.8
2.6
2.4
2.0 ‘
F%
22.22 33.33
37.5
42.86
38.46
41.67
40.00
-
Nature of primary constriction
-
Nearly submedian
Nearly median
Remarks
Chromosome pair with secondary constriction.
-
- -
-
-
~- - Fig . 52b
I Fig. 53b
-- - -
F i g . 54b F ig . 55b
Scale: - % P
Fig. 52b - Leucas stricta Benth. - mitotic metaphase (2n - 22); 53b - E. v e s t i t a Benth. - mitotic metaphase (2n-22); 54b - Leonotis nepetifolia (L.) Ait.f. - mitotic metaphase (2n=28); 55b - Teucrium plectranthoidee Gamble - mitotic metaphase (2n = 32)
Teucrium plectranthoides Gamble (2n=4x=32=A4C2D26)
Normal somatic chromosome number
Chromosome pair with secondary constriction :
Range of chromosome length
Average chromosome length
Total chromatin length
Chromatin length of basic complement
Disparity index (DI)
Variation coefficient (VC)
TF value (%)
2n = 32 (Figs. 55b, 55c)
2
2.2 p to 1.2 p
1.55 p
49.6 ,u
12.4 p
29.41
16.13
42.27
Table 55 --
Detailed Karyotype Analysis (Pig. 5 M )
Chromo- some type
A
A
C
D
D
No. of pairs
1
1
1
4
8
Total length in p
2.2
1.8
2.0
1.6
1.4
D , 1 , 1.2 , 0.4 33.33 Nearly submedian
F%
27.27
22.22 33.33
40.00
37.50
42.86
Short arm length in p
0.6 0.6
0 .4 0 .6
0 . 8
0.6
0.6
-
Nature of primary constriction
-
-
Nearly median
Nearly submedian
Nearly median
Remarks
Chromosome pair with secondary constriction
-
-
-
Fig. 52c
Fig. 54c
Fig. 53c
Fig. 55c
Fig. 52d L-A
Fig. 53d , 4 48 4, 4 , I J I4 14 I,,
Fig. 54d
___JV
A C D
~ i g . 55d $1 $0, fi 11 81 t8 # f #8 #I 88 11 88 11 81 I t A C D
Fig. 52c, 52d - Karyotype and idiogram of Leucas stricta Benth. (211~22) x
2400 approx.; 53c, 53d - Karyotype and idiogram of L.vestita Benth. (2n=22) x 2400 npprox. ; 54c, 54d - karyotype and idiogram of -~eonotis nepetifolia ( L . ) Ait.f. (2n=28) x 2400 approx.; 55c, 55d - Teucrium plectranthoides Gamble (2n=32) x 2400 approx.
Table 56 -- sl l rary of the karymrphetrical analysis on the fifty four e b e r s of South Indian Lariaceae investigated
S1. Name of the taxa 2n n PL KF CPSC RCL ACL TCL CLBC DI VC TF% No. in p in f i in p in p
-- -
Ocimum adscendens Willd.
0. americanum L. - 0. basilicum L.var. - glabratum Benth.
0. basilicum L.var. pilosum Benth.
0. basilicum L.var. - purpurascens Benth.
0. basilicum L.var. - thyrsiflorum Benth.
0. gratissimum L. -
0. gratissimum L.var. - suavis Hook. f . 0. tenuiflorum L.f. - cv. green
0. tenuiflorum L.f. - cv. purple
0. tenuiflorum L.f. - cv. purple-green
S1. Name of the taxa 2n n PL KF CPSC RCL ACL TCL CLBC DI VC TF% No. in p in p in p in pl
12. 0. tenuiflorum L.f. var. hirsuta H0ok.f.
13. Acrocephalus capitatus Benth.
14. Orthosiphon glabratus Benth.
15. - 0. glabratus var. parviflorus Benth.
16. - 0. grandiflorus Bold cv. lilac -
17. - 0. grandiflorus Bold cv. white
18. 0- pallidus Royle
19. - 0. thymiflorus Roth.
20. Plectranthus nummularius Briq.
21. - P. wightii Benth.
22. Coleus aromaticus Benth.
23. - C. aromaticus Benth. var. variegata
S1. Name of the taxa 2n n PL KF CPSC RCL ACL TCL CLBC D I V C T F % No. in &I in p in p in p
2 4 . C. blumei Benth. -- var. verschaffeltii
25. - C. forskohlii Briq.
26. C. laciniatus Benth. - 27. - C. parviflorus Benth.
28. - C. rehneltianus Berger
2 9 . - C. zeylanicus(Benth.) Cramer
30. Anisochilus carnosus (L.f.1 Wall.
31. & eriocephalus Benth.
32. Hyptis capitata Jacq.
33. - H . suaveolens (L.) Poit.
3 4 . Pogostemon benghalensis (Burm.f.) Kuntze
35. - P. heyneanus Benth.
36. & purpurascens Dalz.
37. P, vestitus Benth.
S1. Name of the taxa 2n n PL KF CPSC RCL ACL TCL CLBC D I V C T F % No. in p in p in p in p
38. Eusteralis quadrifolia - (~enth.) Panigrahi 30 - 2x A4D26 2 1.6-1.0 1.16 34.8 17.4 23.08 16.89 38.67
39. Mentha arvensis L. 96 - 8x A4D92 2 1.8-1.0 1.17 112.4 14.05 28.57 17.79 37.98
40. M. rotundifolia (L. ) Huds. 24 - 2x A4D20 - 2 1.6-1.0 1.12 26.8 13.4 23.08 17 .03 41.15
41. M. spicata (L.) Huds. 48 - 4x A4D44 2 1.8-1.0 1.08 51.6 12.9 28.57 16.75 40.53
42. Salvia coccinea Juss. 22 11 2x A2C8D12 1 2.6-1.6 1 .95 42.8 21.4 23.81 18.08 37.92
43. S. leucantha Cav. - 22 11 2x A2C18D2 1 2.8-1.6 2.47 54.4 27.2 27.27 14.35 40.95
44. - S. plebeia R.Br . 16+ 8+ 2x A2C4D10 1 2.6-1.4 1.83 - 29.2 14 .6 30.00 19.27 40.32 0-1B 0-1B 00 u-
45. - S. splendens ker- awl. 44 22 4x A4C2D38 2 2.2-1.2 1 .63 71.6 17 .9 29.41 17.07 41.19
Anisomeles indica (L.) Kuntze
47. - A. malabarica R.Br. 32 - 2x A4D28 2 2.0-1.0 1.36 43.6 21.8 33.33 23.38 41.19
48. Leucas aspera Spreng. 22 - 2x A4D18 2 2.4-1.0 1.44 31.6 15.8 41.18 26.36 37.48 - 49. - L. cephalotes Spreng. 22 - 2x A4D18 2 2.4-1.0 1.47 32.4 16.2 41.18 26.76 38.53
50. 4, linifolia Spreng. 22 - 2x A4D18 2 1.6-1.0 1.27 28.0 14.0 23.08 16.82 37 .95
51. & stricta Benth. 22 - 2x A4D18 2 2.0-1.0 1.35 29.6 14.8 33.33 21.04 37.09
S1. Name of the taxa 2n n PL KF CPSC RCL ACL TCL CLBC D. I. V.C . T.F.% No. in p in in p in p
52. - L. vestita Benth. 22 - 2x A2ClODlO 1 2;4-1.4 1.95 42.8 21.4 26.32 14.56 43.64
53. Leonotis nepetifolia (L.) Ait.f. 28 . - 4x A2C24D2 1 3.6-1.6 2.51 70.4 17.6 38.46 18.98 40.99
54. Teucrium plectranthoides Gamble 32 - 4x A4C2D.26 2 2.2-1.2 1.55 49.6 12.4 29.41 16.13 42.27
Abbreviations used: PL - Ploidy level; K I : Karyotype formula; CPSC - Chromosome pair with secondary constriction; RCL - Range of chromosome length; ACL - Average chromosome length; TCL - Total chromatin length; CLBC - Chromatin length of basic complement; DI - Disparity index; VC - Variation coefficient; TF% - Total forma % (mean centromeric index)
Table 57
Previous and present chromosome counts on different members of the genera investigated in Jimiaceae. The arrangement upto the generic level is essentially based on the corrected version of Bentham's (1876) classification (Sanders and Cantino 1984) 1
Taxon
Tribe - Ocimeae
Sub tribe - Plectranthinae Genus - Ocimum L. 0 . adscendens Willd. -
0. americanum L. -
0 . basilicum L. - ----
Author/~
Sanjappa
Cherian & Kuriachan
Krishnappa & Basavaraj
Saggoo & Bir
Bir & Saggoo
Pushpangadan, et al.
Vij & Kashyap
Singh
Pushpangadan & Sobti
Singh & Sharma
Sobti & Pushpangadan
Singh
Vaarama
Morton
Count Dip- loid num- ber (2n)
-
22
22
-
-
72
' 6 4
84
72
72, 84
72
72, 84
48
48
Year/s
1979
1981
1982
1982, 1983
1985
1975
1975
1980
1982
1982, 1983
1982
1985
1947a
1962
Previous
Hap- loid num- ber (n)
32
-
-
11
11
-
-
-
-
-
-
-
-
-
Present Count
Dip- loid num- ber (2n)
-----
-
-
-
-
-
22
-
-
-
-
-
-
-
72
-
-
Hap- loid num- ber (n)
-
-
-
-
-
11
-
-
-
-
-
-
-
36
-
-
var . " glabratum Benth.
s. var. difforme
s . var. purpurascens var. minima
var. minimum
var. pilosum Benth.
var. purpurascens Benth.
var. thyrsiflora
Sz - Borsos
Mehra & GI11
San jappa
Pushpangadan & Sobti
Sobti & Pushpangadan
Saggoo & Bir
Singh & Sharma
Gill
Ma, et al.
Vembu
Bir & Saggoo
Singh
Pushpangadan, et al.
Singh
Vaarama
Pushpangadan, et al.
Vaarama
Cherian & Kuriachan
Singh
Pushpangadan, et al.
Pushpangadan, et al.
1982
1992
1962
1970
1971
1975
1976b
1979
1981
1981b
1982a
1982
1982
1982
1982
1983
1983
var. thyrsiflorum Benth.
0. campechianum Miller -
0. canum Sins --
-
48
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
52
54
24
22- 32, 34
24, 2 6
- -
24, 26
22
24, 26
2 4 , 26
-
24, 56
-
24
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
6, 11- 14, 17 42
-
-
32+ 0- 4B 32
40
-
40+ 0- 1 B
-
-
-
13
13, 40+ 0- 1 B
-
Krishnappa & Basavaraj
Harley & Heywood
Morton
Sharma
Pal
Pushpangadan, et al.
Vij & Kashyap
Sanjappa
Saggoo & Bir
Singh & Sharma
Bir & Saggoo
Krishnappa & Basavaraj
Pushpangadan & Sobti
Sobti & Pushpangadan
Saggoo & Bir
Singh & Sharma
0. carnosum Link et Otto -
0. gratissimum L. -
embu
ir & Saggoo
ingh
aggoo & Bir
,arley & Heywood
'ushpangadan, et al.
iobti & pushpangadan
jingh & Sharma
aosla & Sobti
Zhosla
rischler
Morton
Mitra & Datta
Pal
Bhattacharya
Singh
Singh & Sharma
Sobti & Pushpangadan
Vembu
Khosla, et al.
Khosla & Sobti
var. suavis Hook. f.
0 . irvinei Mort. - 0 . kilimandscharicum Guerke -
0. nicranthum - 0 . selloi Benth. -
0 . suave Willd. --
Singh
Singh & Sharma
Khosla
Morton
Choudhury, et al.
Kumar, et al.
Bose & Choudhury
Mehra & Gill
' Pushpangadan, et al.
Sobti & Pushpangadan
Saggoo & Bir
Singh & Sharma
I Gill
Bir & Saggoo
Singh
I Sobti & Pushpangadan
I Harley & Heywood
de Wet
I Morton
, Sobti & Pushpangadan
I Renard, et al. / Khosla & Sobti
1 Khosla
0. v i r i d e Willd. -
Genus - Acrocepbalus Benth.
A . c a p i t a t u s Benth. -
Genus - Orthosiphon Benth.
0. d i f f u s e s Benth. -
0. glabratum Benth. -
0. g l ab ra tus Benth. -
var. pa rv i f lo rus Benth.
0. g rand i f lo rus Bold -
cv. lilac
;ingh
'ushpangadan, e t a l .
jobti & Pushpangadan
jingh & Sharma
(hosla, e t a l .
(hosla & Sobti
(hosla
:herian & Kuriachan
Krishnappa & Basavaraj
Cherian & Kuriachan
Krishnappa & Basavaraj
Saggoo & B i r
B i r & Saggoo
Basavaraj & Krishnappa
Krishnappa & Basavaraj
cv. white
0 . incisus Chev. -
0 . incurvus Benth. -
0 . pallidus Royle -
0 . rubicundus (Don) Benth. -
0 . scapiger Benth. - 0. stamineus Benth. -
0 . suffrutescens (Thoun. ) - - Mort.
0 . thymiflorus Roth. -
0 . tomentosus Benth. -
var. tomentosa Hook. -
0 . tubiformis Good --
28
26
28
24
28
28
22
22
-
-
-
28
26
26
48
28
-
-
-
-
-
28
-
-
-
-
-
-
-
-
14
14
14
-
-
-
-
-
14
13
13
13
13
-
28
-
-
-
-
-
-
-
-
-
-
-
28
-
-
-
-
- -
24
-
-
-
-
-
14
-
-
-
-
-
-
-
-
-
-
-
14
-
-
-
-
- -
12
-
-
-
-
-
Morton
Kundu and Sharma
Morton
Chopde
Mehra & Gill
Bhatt
Vembu
Vembu & Sampathkumar
Saggoo & Bir
Gill
Bir & Saggoo
Morton
Krishnappa & Basavaraj
Kundu and Sharma
Cherian & Kuriachan
Morton
Rao & Mwasumbi
Saggoo & Bir
Bir & Saggoo
Saggoo & Bir
Bir & Saggoo
de Wet
1962
198&
1962
1965
1968b, 1972
1974, 1976
1980
1980
1983b
1984
1985
1962
1982
1988a
1984
1962
1981
1982
1985
1983
1985
1958b
0. viscosus Benth. -
Genus - Plectranthus L'Herit.
P. amicorum Blake -
P. argentatus Blake -
P. assurgens (Bak.) Mort. -
P. barbatus -
P. calycinus Benth. -
P. ciliatus Mey. et Benth. - -
P. coesta Buch.- Ham. -
var. macraei Hook.
P. dolichopodus Briq. -
P. dregei Codd -
P. ecklonii Benth. -
P. elegantulus Briq. -
P. esculentus Brown -
P. fruiticosus L'Herit. -
P. gerardianus Benth. -
var. graciliflorus Hook.
P. glandulosus Hook. f. -
P. grallatus Briq. -
P. grandidentatus Guerke -
P. halli Mort. -- P. harrissi Mort. - P. hirtus Benth. --
P. incanus Link - P. insignis Hook. f. - P. japonicus Koidz. -
P. kamerunensis Guerke -
P. laxiflorus Benth. - -
P. macraei Benth. -
P. maddenii Benth. -
P. mollis (Ait.) Spreng. --
P. nilghiricus Benth. -
P. nummularius Briq. -
P. oertendahlii Fries -
P. pachyphyllus Guerke et - Cooke
P. parviflorus Willd. -
P. peglerae Cooke -
P. petiolaris Mey. et Benth. -
P. punctatus L'Herit. -
P. purpuratus Harv. -
de Wet
Morton
Saggoo & Bir
Bir & Saggoo
Morton
Saggoo & Bir
Bir & Saggoo
Cherian & Kuriachan
Saggoo & Bir
Bir & Saggoo
de Wet
Morton
Morton
de Wet
Morton
Hendersson
de Wet
Morton
de Wet
Morton
P. rehmannii Guerke -
P. repens Wall. -
P. rugosus Wall. -
P. saccatus Benth. -
P. scrophularioides Wall. -
P. spectabilis Blake -
P. spicatus May.et Benth. -
P. stocksii Hook. f. -
P. striatus Benth. -
P. strigosus Benth. -
28
28
-
24
24
-
-
28
28
-
-
28
28
-
24
-
24
-
28
( 2 8
-
1 2
-
-
12
12
- -
12
17
-
-
12
-
12
-
12
-
-
P. succulentus Dyer et Bruce -
P. suluensis Cooke - --
-
-
-
28
28
28
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
- -
-
-
-
-
-
-
-
-
-
-
-
- -
-
-
deWet
Morton
Saggoo & Bir
Mehra & Gill
Vij & Kashyap
Bir & Saggoo
Gill
deWet
Morton
Saggoo & Bir
Hendersson
deWet
Morton
Saggoo & Bir
Mehra & gill
Bir & Saggoo
Krishnappa & Basavaraj
Gill
deWet
Morton
-
-
-
deWet
Morton
deWet
p. tenuicaulis(Hook. f.)Mort. -
P. ternifolius Don -
P. thunbergii Benth. -
P. tomentosus Benth. -
P. villosus Cooke -
P. wightii Benth. -
P. woodii Guerke -
Genus - Coleus Lour. C. amboinicus Lour. -
C. aromaticus Benth. -
3hra & Gill
L11
e Wet
or t on
e Wet
orton
e Wet
.orton
:a japathy
kishnappa & Basavaraj
laggoo & Bir
lir & Saggoo
le Wet
(orton
samachandran
Mehra & Gill
Krishnappa & Basavara:
Gill
Scheel
Reddy
Morton
Saggoo & Bir
var. variegata
C. barbatus Benth. -
C. blumei Benth. -
var. verschaffeltii
C. carnosus Hassk. -
C. comosus Hochst. -
C. forskohlii Briq. -
C. frederici Taylor - C. hybridus Hort. -
C. laciniatus Benth. -
C. lanuginosus Hochst. - C. malabaricus Benth. -
C. parviflorus Benth. -
C. pentheri Guerke -
de Wet
Morton
Scheel
Reddy
Morton
Borgnann
Morton
Cherian & Kuriachan
de Wet Morton
Cherian & Kuriachan
Bir & Saggoo
Krishnappa & Basavaraj
Saggoo & Bir
Cherian & Kuriachan
Saggoo & Bir
Cherian & Kuriachan
H. alata (L.f.) Shinners - - s.sp. rugosula (Briq.)
Harley
H. albida Kunth. -
H. argyrophylla Harley -
H. atrorubens Poit. -
H. brevipes Poit. - H. capitata Jacq. -
H. colombiana Epling -
H. conferta Pohl et Benth. -
var. angustifolia Benth.
H. cuneata Pohl et Benth. - --
H. cuniloides Epling - H. emoryi Torr. -
H. eriocephala Benth. -
H. fasciculata Benth. -
H. floribunda Briq. - H. fruticosa Salzm. et Benth. -
H. homalophylla Pohl et - Benth. I
H. inodora Sehrank -
H. irwinii Harley -
H. lanceolata Poir. -
Harley & Heywood
Morton
Harley & Heywood
Chuang, et al.
Reinhard
Harley & Heywood
Harley & Heywood
Baker & Parfitt
Harley & Heywood
Coleman
Harley & Heywood
Morton
H. lantanifolia Poir. -
H. lappulacea Mart. et Benth. - I H. leptostachys -
s.sp. caatingae Harley 1 H. lewocephala Mart. e t -
Benth.
H. lorentziana Hoffm. -
H. macrocephala Mart. et Gal. -
H. macrostachys Benth. - H. martiussi Benth. -
H. microphylla Pohl et Benth. - H. mutabilis (Rich.) Briq. -
H. nudicaulis Benth. -
H. oblongifolia Benth. -
H. pachycephala Epling - H. paludoza St. Hil.et Benth. - H. parkeri Benth. - H. pectinata (L. ) Poit. -
H. pinheiroi Harley -
H.~latanifolia Mart.et Benth. -
H. propinqua Epling - I H. racemulosa Mart. et Benth. - I H. rhomboidea Mart. et Gal. - I H. selloi Benth. - - I H. sidifolia (L'Herit. )Briq. -
H. silvinae Harley -
H..sinuata Pohl et Benth. - I H. spicigera Lam. -
H. stachydifolia Epling - H. suaveolens (L.) Poit. -
Diers
Harley & Heywood 1 1992
Harley & Heywood
Morton
Harley & Heywood
Miege
Morton
Harvey
Pal
Gill & Abubaker
Vij & Kashyap
Bir & Saggoo
Coleman
Krishnappa & Basavaraj
Saggoo & Bir
Vembu
1986
1992
1992
1986
1992
1992
1963
1967
1984
1985
1981
1984
1981
1982
1983
1984
-
-
-
-
-
-
-
-
-
- - -
-
-
-
- -
- -
H. subtilis Epling -
H. tomentosa Poit. -
H. umbrosa Saz: et Benth. - H. urticoides H.B.K. -
H. villicaulis Epling - H. villosa Pohl et Benth. -
Tribe - Uentheae Sub tribe - Pogostemoninae Genus - Pogosteu~n Desf.
(= Dysophylla Blume)
P. auricularia Blume -
P.benghalensis (Burm.f.) - kuntze
P. cablin Benth. -
P. sardneri Hook. -
P. heyneanus Benth. -
Vembu & Ayyangar
Harley & Heywood
Harley & Heywood
Sundberg & Dillon
Harley & Heywood
Chuang, et al.
Hsu
Cherian & Kuriachan
Bir & Saggoo
Lavania
Cherian & Kuriachan
Krishnappa & Basavaraj
Basavaraj & Krishnappa
Cherian & Kuriachan
----
-
14
16
16
8
20
-
-
-
- 6 -
-
-
-
16
16
32
16
16
-
24
28
32
-
16
40
63
90, 96
34
34
34
32
32
-
64
-
-
64
-
-
28
-
- - -
-
-
-
- -
-
64
-
-
- -
- -
64
P. rnollis -
P. paludosus Benth. -
P. paniculatus Benth. -
P. parviflorus Benth. -
P. plectranthoides Desf. -
"
P. pubescens Benth. -
P. purpurascens Dalz. -
P. speciosus Benth. -
P. verticillata Benth. -
P. vestitus Benth. -
P. wightii Benth. -
-
-
- -
32
32
64
64
-
- -
64
-
-
-
-
-
32
34
34
34
72
-
----
16
17
17
16
- -
- -
32
17
32
-
32
16
16
16
16
-
-
-
-
-
16
-
-
-
-
- - - -
-
- -
-
-
-
-
-
- - -
32
-
-
-
32
-
-
-
-
-
-
-
- -
-
- -
- -
-
-
-
- - -
-
-
-
-
Cherian & Kuriachan
Saggoo & Bir
Bir & Saggoo
Cherian & Kuriachan
Krishnappa & Basavaraj
Basavaraj & Krishnappa
Mehra & Gill
Gill
Gill
Mehra & Gill
Gill
Pal
Vij & Kashyap
Saggoo & Bir
Bir & Saggoo
Saggoo & Bir
Krishnappa & Basavaraj
Cherian & Kuriachan
Krishnappa & Basavaraj
Basavaraj & Krishnappa
Borgmann
Cherian & Kuriachan
1981
1982
1985
1981
1982
1983
1968 b
1971 a
1984
1968 b
1971 a, 1984
1971
1975, 1976 b
1982, 1983
1985
1981, 1982a
1981
1982
1984
1982
1983
1964
1981
Genus - Eusteralis Rafin. E. quadrifolia (Benth. ) -
Panigrahi
Sub tribe - Origaninae Genus - Mentha L.
M. alopecuroides Hull. -
M. aquatica L. -
-
-
64
36
-
36
96
96
96
96
96
96
96
96
96
96
96
60, 96
96
96
---- 17
17
-
-
18
-
- - -
-
-
- -
-
-
-
-
- -
-
-
-
-
30
-
- - - - -
-
-
- - -
-
-
-
-
-
-
-
-
-
-
- - - - -
- - - -
-
-
-
-
-
-
-
Krishnappa & Basavaraj
Basavaraj & Krishnappa
Cherian & Kuriachan
Morton
Gill
Schurhoff
Ruttle
Tischler
June11
Rohweder
Graham
Love & Love
Morton
Murray
Gadella & Kliphuis
Olsson
Ouweneel
Ikeda & Ono
Harley & Brighton
Van Loon & de Jong
1982
1983
1984
1956 a
1981 b
1929 b
1931 a
1934
1937
1937
1954
1956 b
1956 a
1958
1963,
1968 b
1967
1968
1969
1977
1978
M. arvensis L. -
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
- -
-
96 +lo5 -
96
96
96
72
72
72, 96
12, 60- 62, 7 2
72
64, 90, 92
72
90
72, 84, 96
72, 96
72
24, 72, 90
36, 92
96
72
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
- -
-
----
-
-
-
-
-
-
- -
- -
-
-
-
-
- -
- -
-
Tucker, et al.
Roy,etal.
Fernandes & Leitao
Singh
Schurhoff
Lietz
Ruttle
Junell
Rohweder
Nagao
Morton
Sharma & Bhattacharya
Ikeda & Seiroku
Ikeda & Udo
Olsson
Ouweneel
Taylor & Mulligan
Belyaeva & Kovineva
Harley & Brighton
1980
1983
1984
1984, 1985
1929 b
1930
1931 a
1937
1937
1941 a
1956 a
1959 a
1966
1967
1967
1968
1968
1972
1977
1980
1981 b
1984
1985
1985
1986
1990
1977
1982 a
1971 b, 1975 1986
1965
1985
1965
1972
1972
1929 b
1929
1980
Tucker, et al.
Gill
Fernandes & Leitao
Kundu & Sharma
Parfenov & Dimitrieva
Pogan, et al.
Tucker & Fair brothers
Harley & Brighton
Love & Love
Sobti
Tyagi
Sobti
Kundu & Sharma
Sobti
Belyaeva & Kovineva
Ono
Schurhoff
Wolf
Tucker, et al.
-
- -
-
-
-
-
-
-
- -
-
- -
-
-
-
-
-
-
-
-
-
-
-
-
96
-
-
-
-
-
-
-
- -
-
-
-
---- -
48
-
-
-
- -
-
-
- -
-
-
-
- - -
-
-
f. piperascens Holmes
M. arvensis L. -
s.sp. borealis (Michx.) Taylor et MacBryde
s.sp. haplocalyx var. piperascens
Holm.
M. arvensis - var . javanica (Blume)
H0ok.f.
var. piperascens Holm.
M. canadensis L. -
72
-
24, 72, 90
96
72, 74
36
72
96
72
96
96
72
60
96
96
96
54
54
96
M. cervina L. -
M. citrata - M. cordifolia -
M. crispa L. -
M. cunninghamii Benth. -
M. dumetorum -
M. gattefossei Maine -
M. gentilis L. -
36
36
36
96
3 6 , 48
48
84
4 8 , 84
72
96
40
20
48
+ 32 -
54 , 60 8 4 , 96 ,
108, 120
54
7 2
72
54
54 , 60 8 4 , 96 ,
108 , 120
- -
-
-
-
- -
-
- -
-
-
-
-
-
- - -
-
-
---- -
-
-
-
-
- -
-
-
-
-
-
-
-
-
- - -
-
-
-
-
-
-
-
- -
-
-
-
-
-
- -
-
-
-
-
-
-
Harley & Brighton
Tucker, et al.
Fernandes & Leitao
Sobti
Sobti
Murray
Harley & Brighton
Tucker, et al.
Hair & Beuzenberg
Sobti
Morton
Sobti
Makarov & Reznikova
Harley & Brighton
Morton
Sobti
Baquar
Tucker, et al.
Gill
Fernandes & Leitao
1977
1980
1984
1965
1962 b, 1965
1960 b
1977
1980
1960
1965
1956 a
1965
1972
1977
1956 a
1965
1967
1980
1981 b
1984
M. gracilis Sole. -
M. japonica Makino - M. lamarckii -
M. lavandulidora Sacco -
M. longifolia (L.) Huds. -
-
-
- -
-
- - -
- -
-
- -
-
-
-
-
-
-
-
-
-
54
60 , 7 2 , 8 4 , 96
49
36
96
18
18
24
24
18
48
48
24
36 , 48
24
18, 27
24 36 , 48
36
24
24
24
24
---- -
-
- -
-
- -
- -
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
- -
-
-
-
-
- -
-
-
-
-
-
-
-
Gill
Tucker & Fairbrothers
NagAo
Delay
Shimizu, et al.
Schurhoff
Lietz
Ruttle
June11
Heimans
Nagao
Polya
Tsuda
Morton
Murray
Sobti
Zhukova
Ouweneel
Majovsky, et al.
Sobti
Markowa & Iwanowa
1981 b
1990
1941 a
1947
1967
1929 b
1930
1931 a
1937
1938
1941 a
1950
1954
1956 a
1958, 1960 b
1962 b
1965
1967 b
1968
1970 a
1971 a,b
1972
s.sp. hymalaiensis Briq.
s.sp. lavandulacea Schimp ex Briq.
s.sp. longifolia
s.sp. typhoides Briq.
var. cyprica (Braun) Briq.
M. longifolia (L.) Huds. -
var. grandis Briq.
var. typica Fiori
M. microphylla Koch. -
M. muelleriana Schultz -
M. nemorosa Willd. -
M. niliaca Jacq.et Briq. -
M. parniroalaica Boriss -
M. piperita L. -
Wolf
Ruttle
Nagao
Morton
Murray
Sharma & Bhattacharya
Sobti
Sobti
Baquar
Ouweneel
Belyaeva, et al.
Belyaeva & Kovineva
Bugaenko & Reznikova
Gill
Roy, et al.
Kundu & Sharma
M. sacchalinensis (Briq.) - Kudo
M. sapida -
M. satureioides R. Br. - -
M. scotica Graham - var. lacerata
M. smithian? Graham -
M. spicata (L. ) Huds. -
Belyaeva & Kovineva
Sobti
Harley & Brighton
Morton
Sobti
Ruttle
Nagao
Suzuka & Koriba
Morton
Murray
Sobti
Baquar
Harley & Brighton
Bugaenko & Reznikova
Pucker, et al.
:ill
coy, et al.
?ernandes & Leitao
Cundu & Sharma
Tucker & Fairbrothers
Harley & Brighton
Tucker, et al.
Harley & Brighton
Tucker, et al.
Harley & Brighton
Fernandes & Leitao
Silvestre
Harley & Brighton
Harley & Brighton
Schurhof f
Ruttle
Tischler
Rohweder
Delay
Bir & Saggoo
Gill
Saggoo & Bir
Heimans
Schurhof f
Lietz
var. acutifolia (Sm. ) Fraser
M. villosonervata Opiz. -
M. viridis L. -
var. typica
f. brevipetiolata (Rehb. ) Fiori
42 , 84 ,
120, 132
7 2 , 8 4 , 96,
120
96
7 2 , 7 8 , 84
7 8 , 8 4 , 90,
120, 132
84
48
36
36
32, 48
-
-
48
-
-
- -
-
-
- - -
-
24
24
-
----
---- -
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
- -
-
-
-
-
-
Morton
Baquar & Reese
Sobti
Suoninen
Olsson
VanLoon & de Jong
Morton
Schurhoff
Delay
Sharma & Bhattacharya
Saggoo & Bir
Bir & Saggoo
Sacco, et al.
1956 a
1965
1965
1966
1967
1978
1956 a
1929 b
1947
1959 a
1983
1984
1968
Tribe : Salvieae I Genus : Salvia L. I S. aegyptiaca L. -
S. aethiopis L. -
S. albimaculata Hedge et - Huber - Morath
S. alborosea Epling et Jativa
S. amarissima Ort. -
S. amasiaca Freyn et Bornm. - S. amplexicaulis Lam. -
S. argentea L .
S. armenekensis Rech. -
S. armeniaca -
Delestaing
Bhattacharya, et al.
Humphires, et al.
Markova & Ivanova
Afzal - Rafii Sopova, et al.
Fernandes & Leitao
Singh
Afzal-Rafii
Harley & Heywood
Mercardo, et al.
Aryavand
Markova & Penka
Strid & Franzen
Markova & Ivanova
Markova & Thu
Afzal-Rafii
Strid & Franzen
Markova & Ivanova
Fernandes & Leitao
Afzal-Rafii
Patudin, et al.
S. atropatana Bge. - S.austriaca - S. azurea Lam. -
S. beckeri -
S. brachysiphon Stapf. - S. breviflora Moc. et Sesse -
S. cadmica Boiss - S. caespitosa Montbr. et -
Heldr .
S. campanulata Wall. - var. hirtella Stibal
var. nepalensis Stibal
S. candelatrum -
S. candidissima Vahl -
S. carduacea -
S. castanea Dielz.
S. ceratophylla L. -
S. chloroleuca Rech.f. et - Allen
S. chrysophylla Stapf. -
S. cilicica Boiss. -
I Afzal-Rafii
1 Markova & Ivanova
Scheel
1 Patudin, et al.
Afzal - Rafii
Harley & Heywood
Afzal - Rafii
Saggoo & Bir
Rosua & Gabriel
Van Loon & Snelders
Afzal - Rafii
Stewart
Epling, et al.
Gill
Afzal - Rafii
Fernandes & Leitao
Haque
Sugiura
Hruby
Delestaing
Erbrich
Harvey
Bhattacharya
Haque 1 1981
Krishnappa & Basavaraj 1982 I Saggoo & Bir
Gill
Bir & Saggoo
Harley & Heywood
Haque & Ghoshal I lg80a
Scheel
Gill
Bir & Saggoo
Gill
S. columbariae -
S. columbariae -
S. crassifolia Sibth. et Sm. -
S. cryptantha Montbr.et Auch - S. deserta - S. desoleana Atzei et Picci - S. discolor Kunth - S. dorisiana Fernald -
S. dumetorum Andrz. -
S. elegans Vahl. - var. sonorensis Fernald I
S. farinacea Benth. -
cv. " Lavender Blue'
cv. 'Royal Blue' I
cv. 'White'
Haque & Ghoshal
Stewart
Epling, et al.
Afzal - Rafii
Diana - Corrias
Harley & Heywood
Sugiura
Delestaing
Harley & Heywood
Sugiura
Gill
Bir, et al.
Gill
Haque
Haque & Ghoshal
S. fluviatilis Fern. -
S. forskohlei L. -
S. frigida Boiss.
S. fruticosa Mill. -
S . gerneriflora Lindl - Paxton
S. glutinosa L. -
S. grahamii Benth. -
Mercardo, et al.
Afzal-Rafii
Markova & Penka
Markova & Ivanova
Afzal-Rafii
Harley & Heywood
Scheel
Hruby
Polya
Linnert
Majovsky, et al.
Markova & Ivanova
Skalinska, et al.
Afzal-Rafii
Haque & Ghoshal
Strid & Franzen
Love & Love
Markova & Ivanova
Singh
Bir & Saggoo
Gill
Raman & Kesavan
Haque & Ghoshal
Haque
Haque
Saggoo & Bir
Bir & Saggoo
Afzal - Rafii
Markova & Ivanova
Palomino, et al.
Harley & Heywood
Ward
Scheel
Gill
Vir-Jee & Kachroo
Palomino, et al.
Afzal-Rafii
Haque & Ghoshal
Haque
Mercardo, et al.
Estilai, et al.
Linnert
Afzal-Rafii
Haque & Ghoshal
S. hydrangea DC ex Benth. - S. hypargeia Fisch. et Mey. - S. hypoleuca Benth. -
S. interrupta - s.sp. pani
S. involucrata -
S. itatiaiensis Dusem - S. judaica Boiss.
S. jurisicii Kosanin - .-
S. lanata Roxb. -
S. lanigera Poir. -
S. lavandulifolia Vahl -
S. lavandulifolia Vahl -- s.sp. amethystea
s.sp. blancoana
s.sp. maurorum
s.sp. mesatlantica
s. sp. Eyrenaeorum
s.sp. vellerea
s.sp. vellerea var. lagascana
Sopova, et al.
Afzal-Rafii
Aryavand
Afzal-Rafii
Rosua & Gabriel
Gill
Harley & Heywood
Afzal-Rafii
Sopova, et al.
Mehra & Gill
Gill
Sarkar, et al.
Kliphuis
Ubera
Rosua & Gabriel
S. lavanduloides Kunth
S. leucantha Cav.
S. limbata Mey. -
S. longispicata Mart. et -
S. lupulina Fern. -
S. macilenta Boiss. -
S.macrosiphon Boiss. -
S. merjamie Forsk. -
S. mexicana L. -
S. microphylla Kunth -
S. microstegia Boiss. et -
Palomino, et al.
Carlson & Stuart
Gill
Vij & Kashyap
Bhattacharya
Haque & Ghoshal
Bir & Saggoo
Haque
Gill
Afzal - Rafii
Mercardo, et al.
Palomino, et al.
Afzal-Rafii
Delestaing
Aryavand
Hedberg & Hedberg
Palomino, et al.
Harley & Heywood
Afzal-Rafii
S. molesta Mitchell -
S. montbretti -
S. moorcroftiana Wall. -
S. multicaulis Vahl
S. munzii Epling -
S. napifolia Jacq. -
S. nemorosa L. -
s. sp. nemorosa
I Kuriachan
/ Bhatt, et al. 1 Bir & Saggoo
Gill I Aryavand
Afzal-Rafii
Baker & Parfitt
Harley & Heywood
Afzal-Rafii
Hruby
Benoist
Tischler
Linnert
Chauhan & Abel
Majovsky, et al.
Aryavand
Afzal-Rafii
Afzal-Rafii
Markova & Ivanova
Markova & Penka
Van Loon & de Jong
S. nemorosa L. var. pseudo-sylvestris (Stapf.) Bornm.
S. nilotica Juss. et Jacq. - S. nubicola Benth. -
S. nutans - S. occidentalis Sw. -
S. ochrantha Epling - S. officinalis L. -
S. ombrophila Dusen - S. ~alestina Benth. -
S. persepolitana Boiss. -
14
32
12- 24
16 , 18 , 20 , 22, 24
22
18
32
14
14
14
14
14
14
14
14
14
14
-
240
20
-
-
-
-
-
-
-
-
-
-
- -
- -
-
- 7 -
- -
-
7
- -
10
-
-
-
-
-
-
-
- -
-
-
-
-
-
-
-
-
- - -
-
-
-
-
-
- -
- -
-
-
-
-
-
-
-
-
-
-
-
- -
-
Afzal - Rafii
Hedberg & Hedberg
Pal
Bhattacharya
Markova & Ivanova
Harley & Heywood
Scheel
Hruby
Suzuka & Koriba
Linnert
Butterfass
Love & Love
Gill
Af zal-Rafii
Haque & Ghoshal
Van Loon
Fernandes & Leitao
Gill
Harley & Heywood
Af zal-Rafii
Aryavand
S. phlomoides Asso. -
S. pinnata L. -
S. pisidica Boiss. et Heldr. -
S. plebeia R.Br. -
S. polystachya Ort. -
S. pomifera L. - s. sp. calycina (Sm. )
Hayek
s.sp. pomifera
S. potentillaefolia Boiss.et - Heldr .
S. pratensis L. -
S. pseudococcinea Jacq. -
20
22
14
16
16
16+ 1 B
16
-
22
14
14
14
16
16
18
18
18
18
18, 32
-
-
-
-
-
-
-
-
8+ 1 B
-
- -
-
- -
-
-
-
- 8 -
-
-
11
-
-
-
-
-
-
-
-
16+ 0-1B
-
- -
-
-
-
- -
-
-
-
-
-
-
-
-
-
-
-
-
8+ 0-1B
-
-
- -
-
-
-
-
-
-
-
-
-
Afzal - Rafii
Chauhan & Abel
Markova & Cerneva
Af zal-Rafii
Mehra & Gill
Gill
Vij & Kashyap
Gill
Harley & Heywood
Von Bother
Afzal - Rafii
Van Loon
Frey, et al.
Haque
Love & Love
Markova & Ivanova
Gill
1981
1968
1982, 1984
1980
1968b
1971
1975, 19761,
1984
1992
1970a
1980
1971
1980
1980
1981
1982
1982b
1982a
1984
S. purpurea Cav. -
S. guerceto-pinorum Epling - S. recognita Fisch. et Mey. -
S. relexa Hornem. -
S. reuteriana Boiss. - S. rhyacophila Epling -
S. ringens Sibth. et Smith -
var. macedonica Briq.
S. russegeri Fenzl. -
S. rutilans Carr. - S. sagittata R. & P. - S. sahandica Boiss. et Buhse -
S . scabiosifolia Lam. -
S. sclarea L. -
S. sclareoides Brot. -
S. sessei Benth. -
S. silvarum Epling -
22
22
16
22
20
-
-
12
16
12
14, 16
20
18
22
22
14
22
22
22
22
22
22
20
22
66
-
- - -
10
10
10
6
-
- -
-
9
11
-
- - -
- -
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
- - - -
-
-
-
-
-
-
-
-
-
-
- - - -
-
-
-
-
-
-
- -
-
-
- - -
-
-
-
-
- - -
Harley & Heywood
Afzal-Rafii
Haque & Ghoshal
Haque
Ghaffari
Harley & Heywood
Af zal-Raf ii
Markova & Ivanova
Af zal-Raf ii
Raman & Kesavan
Harley & Heywood
Aryavand
Afzal - Rafii Markova & Ivanova
Fujita
Afzal - Rafii
Van Loon & de Jong
Diana - Corrias Fernandes & Leitao
Elnir, et al.
Fernandes & Leitao
Harley & Heywood
S. tomentella Pohl -
S. tomentosa -
S. tonolea Boiss. - S. triloba L.fil -
S. urica Epling --
S. urnapuana Epling -
S. verbascifolia -
S. verbenacea L. -
22
-
22
22
86
-
22
14
14
22
12
16
64
54
42
59
4 2 , 5 4 , 5 8 , 6 0 , 64
54
4 8 , 5 4 , 58- 6 0 , 64
64
42
-
11
-
-
-
7. 8
- - -
-
-
- - - -
-
-
- -
-
-
-
-
-
-
- -
- - - -
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
- -
- -
- -
- - -
-
-
-
-
-
Haque
Mercardo, et al.
Harley & Heywood
Afzal - Rafii
Ferrarella, et al.
Fernandes & Leitao
Harley & Heywood
Palomino, et a1 .
Af zal-Rafii
Yakovleva
Benoist
Reese
Gadella & Kliphuis
Bjorkqvist, et al.
Bhattacharya, et al.
Dahlgren, et al.
Van Loon, et al.
Kramer, et al.
S. verbenacea L. -
s.sp. multifida
S. verticillata L. -
S. virgata Ait. -
S. viridis L. -
S. viridis L. -
S. viscosa Jacq. -
S. widemanni Boiss. - S. xanthocheila Boiss. - Tribe: Lamieae
Subtribe: Laminae
Genus : Anisomeles R. Br.
A. heyneana -
A. indica (L.) Kuntze -
A. malabarica R.Br. -
16
16
32 , 40
14
22
34
34
34
30
40
34
34
34
34
34
-
-
-
34
32
-
-
-
-
-
-
-
- -
20
-
-
-
-
-
17
17
17
-
-
-
- -
- -
-
- -
-
-
-
- - - -
-
-
-
-
34
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
Afzal - Rafii
Markova & Ivanova
Afzal - Rafii
Krishnappa & Basavaraj
Mehra & Gill
Gill
Sharma
Bir & Sidhu
Vij & Kashyap
Vembu
Cherian & Kuriachan
Vembu & Sampathkumar
Krishnappa & Basavaraj
Saggoo & Bir
Gill
Bir & Saggoo
Kundu & Sharma
Vembu & Sampathkumar
1980, 1981
1982a
1980
1972
1981
1982
1968b
1970
1970
1974, 1980
1975, 1976b
1979
1981
1981
1982
1983
1984
1985
1988a
1978,
-
- -
-
-
-
22
-
-
-
-
-
- -
-
-
- -
-
- - -
-
-
1968b
1970
1975, 1976b
1982
1984
1991
1982, 1983
1984
1982
1982, 1983
1983
1980
1980
1980
1984
1982
1982, 1983b
1983
1984
1980
1980
1982
1983b
- -
-
-
-
-
-
- - -
-
-
-
- -
-
-
-
-
-
-
-
-
-
-
-
-
14
11
15
14
-
11
-
-
- -
11
-
11
-
11
-
-
-
11
L. 3halotes Spreng. -
L. chinensis -
L. ciliata -
L. clarkei Hook. f. -
L. deflexa Hook. f. -
L. diffusa Benth. -
L. eriostoma Hook. - L. helianthemifolia Desf. -
L. hirta Spreng. --
Mehra & Gill
Gill
Vij & Kashyap
Krishnappa & Basavaraj
Gill
Singh, et al.
Saggoo & Bir
Cherian & Kuriachan
Krishnappa & Basavaraj
Saggoo & Bir
Renard, et al.
Ayyangar & Vembu
Krishnan
Vembu & Sampathkumar
Cherian & Kuriachan
Krishnappa & Basavaraj
Saggoo & Bir
Cherian & Kuriachan
Ayyangar & Vembu
Vembu & Sampathkumar
Krishnappa & Basavaraj
Saggoo & Bir
28
28
22
22
-
-
-
-
22
-
28
22
22
22
-
22
-
22
-
22
22
22
-
L. lamifolia Desf. -
L. lanata Benth. -
L. lanceaefolia Desf. -
L. lavandulaefolia Rees - L. lavandulaefolia Rees -
L. linifolia Spreng. -
L. marrubioides Desf. - L. martinicensis R. Br. -
L. mollissima Wall. -
L. mollissima Wall. - var. scaberula Hook.
-
28
30
-
28
-
22
-
22
-
22
-
-
-
28
28
28
-
- -
11
-
-
14
-
14
-
11
-
11, 11+ 0- 1 B
-
11+ 0- 1 B
14
14
-
-
-
14
14
14
-
-
- -
-
-
-
-
-
-
-
-
22
-
-
-
-
-
-
-
-
138
-
-
-
-
-
-
-
-
-
-
-
-
-
- -
-
- -
- -
Gill
Bhatt, et al.
Bir & Saggoo
Krishnappa & Basavaraj
Gill
Krishnappa & Basavaraj
Saggoo & Bir
Krishnappa & Basavaraj
Saggoo & Bir
Chopde
Saggoo & Bir
Cherian & Kuriachan
- Gill
Krishnappa & Basavaraj
Gill
Krishnappa & Basavaraj
Gill
Saggoo & Bir
Bir & Saggoo
1982, 19831,
1970
1975
1981, 1984
1982
1984
1982
1982, 1983b
1982
1983b
1965
1982
1981
1978
1982
1970
1982
1984
1981
1981, 1982a
L. montana Spreng. -
L. nutans Spreng. -
L. procumbens Desf. -
L. stricta Benth. -
L. suffruticosa Benth. -
L. urticaefolia R. Br. - L. vestita Benth. -
L. zeylanica R. Br. -
Genus - Leonotis R.Br.
var. nepetifolia
L. africana Briq. - L. nepetifolia (L.) Ait.f. -
Tribe - A-jugeae
Genus - Teucrium L.
28
-
15
22
28
-
22
22
-
24
28
-
14
-
-
-
11
-
-
11
T. aragonense Loscos et Pardo -
T. arduini L. -
-
-
-
-
-
22
-
-
-
-
22
-
-
52
32
-
-
-
-
-
-
-
-
-
- -
- -
-
-
Krishnappa & Basavaraj
Bir & Saggoo
Saggoo & Bir
Krishnan
Krishnappa & Basavaraj
Saggoo & Bir
Cherian & Kuriachan
Krishnappa & Basavaraj
Saggoo & Bir
Miege
Morton
-
-
1982
1980, 1981, 1984
1982, 1983
1980
1982
1982, 1983
1984
1982
1982, 1983a
-
-
Puech
June11
T. argutum R. Br. -
T. aureum Schreb. -
s.sp. angustifolium (Willk.)Puech
s.sp. aureum
s.sp. latifolium (Willk.)Puech
T. belion Schreb. -
T. bicolor (Sm.) Rees - T. botrys L. -
T. brevifolium Schreb. -
T. brachyandrum Puech - T. canadense -
s.sp. viscidurn (Piper) Taylor et Macbryde
T. capitatum L. -
var. polycephalum (Pomel) Briq.
T. carthaginense -
T. chamaedrys L. -
32
32
-
26
52
26
26
90
-
10
32
30
7 4
32
26
26 , 52
26
52
52
26
32
64
-
-
- -
- -
-
-
- -
- -
-
- -
-
-
- -
-
- -
- -
16
-
-
-
- -
14
- -
-
-
- -
-
-
- -
-
- -
Delay
Gill
Puech
Carmen
June11
Van Loon & Kieft
Strid
Puech
Love & Love
Puech
Fernandes & Leitao
Puech
Puech
Delay
Reese
- - - -
- -
-
-
-
-
-
- -
-
-
-
-
- - -
-
-
T. haenseleri Boiss. - T. halacscynanum -
T. hervieri Barnades - T. hircanicum -
T. lamifolium D'urv. -
T. lucidum L. -
T. mairei Senn. -
T. massilense L. -
T. mesanidum (Lit. et Maire) -- Sauv. et Vindt.
T. montanum L. -
ssp. montanum
T. nuchense -
I
Fernandes & Leitao
' Damboldt .
, Messia & Rejon
i Koudysheva & Akhmed- Zade
Markova & Cerneva
Fernandez, et al.
Puech
Van Loon & de Jong 1 1978 Contandriopoulos 1 1957c Fernandez, et al. 1 1977
Puech
Guinochet & Logeois
Van Loon & de Jong
Fernandez, et al.
Love & Love
Markova & Cerneva
Afzal-Rafii, et al.
Singh
Baksay
1978
Koudysheva & Akhmed - Zade 1 1968
T. orientale L. -
T. plectranthoides Gamble - I T. polium L. -
s.sp. aguilasense Puech I s.sp. capitatum (L.)
Briq.
var. majoricum (Rouy) Wk.
T. polium L. - s. sp. clape Puech I s.sp. gabesianum Le
Houerou
Aryavand 1977
Ga japathy 1962b
Lorenzo-Andreu & Garcia - Sanz.
Reese
Koudysheva & Akhmed - Zade 1 1968 Gill / 197lb
Markova & Thu I lg7* Puech
Puech
Vij & Kashyap
Puech
Sopova, et al.
Afzal - Rafii, et al.
Puech
Fernandez, et al. 1977
Markova & Cerneva 1982
Guinochet & Le Franc 1972
Puech 1 1911
s.sp. polium
s.sp. purpurascens (Benth.) Puech
s.sp. vincentianum (Rouy) Wood
T. pseudochaemopitys L. -
T. quadrifarium Buch - Ham. -
T. guadrifarium Buch - Ham. -
T. rouyanum -
T. royleanum Wall. -
T. salviastrum Schreber - T. sauvagei Le Houerou -
T. scordium L. - T. scordioides Schreber -
T. scorodonia L. -
T. stoloniferum Roxb. -
T. subspinosum Pourr. -
T. tomentosum Heyne -
T. vincentinum Rouy -
- Markova & Cerneva
- Ubera
- Malla, et al.
- Saggoo & Bir
- Gill
- ] Puech
-
- Gill
- 1 Bir & Saggoo
- Gill
- Fernandes & Leitao
- Puech
- Morton
- Fernandes & Leitao
-
- Bhattacharya
- 1 Guinochet & Lefranc
- Krishnappa & Basavaraj
- Cherian & Kuriachan
- Fernandes & Leitao
D..
m i 01
F i q . 56 C m p a r a t i v e id iog ram p l a t e o f d i f f e r e n t t a x a i n v e s t i g a t e d i n Lamiaceae. (Contd.) 12d - - 0. tenu i f l o ru rn L. f . cv . purp le-green, 13d - & t e n u i f l o r u m L.f. var . hirsuta H0ok.f.. 14d - Acrocephalus c a p i t a t u s Benth., 15d - Orthos iphon g l a b r a t u s Benth., 16d - g. q l a b r a t u s v a r . p a r v i f l o r u s Benth., 17d - 2. q r a n d i f l o r u s Bo ld cv. lilac, 18d - & q r a n d i f l o r u s Bo ld cv. white, 19d - g. p a l l i d u s Royle, 20d - 0. t h y m i f l o r u s Roth., 21d - P l e c t r a n t h u s numrnularius Br iq . , 22d - P. w i g h t i i Benth.
F i g . 56 Comparat ive i d i o g r a m p l a t e o f d i f f e r e n t t a x a i n v e s t i g a t e d i n Lamiaceae. (Contd.) 23d-Coleus a r a n a t i c u s Benth., 24d - i . a romat i cus Benth. var . v a r i e g a t a , 25d-C. bluernei Benth. var . v e r s c h a f f e l t i i , 26d- f o r s k o h l i i Br iq . , 27d-5. l a c i n i a t u s Benth., 28d-C. - e a r v i f l o r u s Benth., 29d- C. r e h n e l t i a n u s Berger, 3Dd-C. z e y l a n i c u s (Benth.) Cramer, 3 l d - A n i s o c h i l u s carnosus (L . f ) Wal l , 32d- - - A.er iocephalus Benth., 3 3 d - r n c a p i t a t a Jacq. -
F ig . 56 Comparat ive id iog ram p l a t e of d i f f e r e n t t a x a i n v e s t i g a t e d i n Lamiaceae. (Contd.)
34d-H. - suaveolens (L.) Po i t . , 35d-.Pagostemon benghalens is (6urm.f.) Kuntze, 36d-E. heyneanus Benth., 37d - P, purpurascens Dalz., 38d-c. v e s t i t u s Benth., 39d-Eus te ra l i s q u a d r i f o l i a (Benth.) Pan ig rah i , 40d - Mentha a rvens is L., 41d- - M. r o t u n d i f o l i a (L.) Huds., 42d - - M. spicata (L.) Huds.
F i g . 56 Compara t i ve i d i o g r a m p l a t e o f d i f f e r e n t t a x a i n v e s t i g a t e d i n Lamiaceae. (Contd.)
43d - c o c c i n e a Juss., 44d - 2. l e u c a n t h a Cav., 45d - 8. p l e b e i a R.Br. , 46d - 2. sp lendens Ke r - Gawl, 47d - Anisomeles indica (L.) Kuntze, 48d- A. m a l a b a r i c a R. Br., 49d-Leucas aspe ra Spreng., 50d- L. c e p h a l o t e s Spreng., 5 l d - L - l i n i f o l i a Spreng., 52d - L. s t r i c t a Benth., 53d - L. v e s t i t a Benth., - 54d - L e o n o t i s n e p e t i f o l i a ( L . ) A i t . f . , 55d- - Teucr ium p l e c t r a n t h o i d e s Gamble
16 18 22 24 26 28 30 32 34 36 40 44 48 64 72 96
CHROMOSOME NUMBERS
Fig. 57 Range of chromosome numbers recognized in the present study
6 w t 8 9 10 11 12 13 15 16 17
BASE NUMBERS
Fig. 58 Range of base numbers recognized i n the present study
2 X 4 X 6 X 8 X PLOIDY LEVEL
Fig. 59 Range of ploidy l e v e l recognized i n the present study
I - annual herb I - perennial herb
80 I11 - annual undershrub - perennial undershrub
7 0
6 0
Q 50 a rl 0 rl
E 40 rl 0 a
30 0
w M
2 20 2 0 $4
2 10
0 I I1 111 IV
GROWH HABIT
Fig. 60 Range of polyploids recognized under different growth habits in the present sutdy
~ig. 62: Comparison of the major karyotypic parameters in the cytotypes of Acrocephalus Benth., Orthosiphon Benth. and Plectranthus L' Herit. TCL - Total chromatin length, CLBC - Chromatin length of basic complement, TF% - Total forma percentage, DI - Disparity index, VI - variation coefficient. 13 - Acrocephalus capitatus Benth. (2n=18), 14 - Orthosiphon glabratus Benth. (2n=26), 15 - 0. &labratus var. parviflorus Benth. (2n =26), 16 - 0. grandiflorus Bold cv. lilac (2n=28), 17 - 0. grandTflorus Bold cv. white (2n=28), 18 - 0. pallidus ~oTle (2n=28), 19 - 0. thymiflorus Roth. (2n=24), 20 - Plectranthus - - -- nummularius Briq. (2n=28), 21 - P. wightii Benth. (2~24)
TCL CLBC 0 rn D1 vc
Fig. 63 Comparison of major karyotypic parameters in the cytotypes of Coleus Lour. TCL - Total chromatin length, CLBC - Chromatin length of basic complement, TF% - Total forma percentage, DI - Disparity index, VC - variation coefficient. 22 - Coleus aromaticus Benth. (2n=32), 23 - C. aromaticus Benth. var. variegata (2n=32), 24 - C. blumei Benth. var. verschaffeltii (2n=48), 25 - C. forskohlii Briq. (2n=28), 26 - C. - lacintatus Benth. (2n=48),arviflorus ~enth; (2n=72), 28 - C. rehneltianus ~ e r ~ e r - (2n=48), 29 - - C. zeylanicus (~entF~ramer (2~28)
Fig. 64 Comparison of major karyotypic parameters in the cytotypes of Anisochilus Wall., Hyptis Jacq. and Pogostemon Desf. TCL - Total chromatin length, CLBC - Chromatin length of basic complement, TF% - Total forma percentage, DI - Disparity index, VC - Variation coefficient. 30 - Anisochilus carnosus (L.f.) Wall. (2n=34), 31 - A. eriocephalus Benth. (2n=32), 32 - Hyptis capitata Jacq. (2n=32), 33 - H. suaveolens (L.T Poit. (2n=28), 34 - Pogostemon benghalensis (Burm.f. ) Kuntze (2n=64), 35 P. heyneanus Benth. (2n=64), 36 - P. - purpurascens Dalz. (2n=32), 37 - - P. vestitus Benth. (2n=32)
TCL CLBC TF% DI vc
Fig. 65 Comparison of the major karyotypic parameters in the cytotypes of Eusteralis Rafin., Mentha L. and Salvia L. TCL - Total chromatin length, CLBC - Chromatin length of basic complement, TF% - Total forma percentage,DI - Disparity index, VC - Variation coefficient. 38 - Eusteralis uadrifolia (Benth.) Panigrahi,(211=30),?9-Mentha arvensis L. (2n=96), 40 - M. rotundifolia L . 2 41 - g. spicata (L.) ~udn==48-- Salvia coccinea .Tun;. (2n=22), 43 - S. leucantha Cav. (2n=22), 3 4 -- S. plebeia R. Br., (2n=16+0-1 B) - - 45 - - S. splendens Ker-Gawl. (2~44)
TCL CLBC 0 TF% vc -
Fig.66 Comparison of the major karyotypic parameters in the cytotypes of Anisomeles R.Br., Leucas R.Br., Leonot R.Br. and Teucrium L. TCL - Total chromatin length, CLBC - Chromatin length of basic complement, TF% - Total for) percentage, DI - Disparity index, VC - Variation coefficient 46 - Anisomeles Indica (L.) Kuntze (2n = 34), 47 - A. malabarica R. Br. (2n = 32), 48 - Leucas aspera Spren, (2n = 22), 49 - L. cephalotes Spreng. (2n = 22), 50 - LT linifolia Spreng. (2n = 221, 51 - L. stricta Benth.(2n=2, 52 - L. vestita xenth. (2n=22), 53 Leonotis nepetifoKa (L.) Ait.f. (2n=28), 54 - ~eucrium plectranthoides Gamb. (2n=3T)
Xi - Primary base number
X2 - Secondary base number
PAP - Proto auto ploidy AP - Amphiploidy AD - Ascending dysploidy
DD - Descending dysploidy
2
Fig. 67 Phylogenetical scheme showing the probable evolution of chromosome numbers in the taxa investigated. (Bold numbers in the square's represent chromosome numbers identified)
DISCUSSION
a. Chromosome data in the genera investigated
Chromosome counts are made on fifty four members of Lamiaceae prevailing
in South India. Cytologically the family is variable. The chromosome
spectrum varies from 2n = 16 + 0-1B to 96, with majority of the species
concentrated in the number 2n = 32, followed by 2n = 48, 2n = 28 and
2n = 22 (vide Table 56; Fig. 57) . The presence of identical numbers in
unrelated genera is a very noteworthy feature in this family (Fedorov,
1969). The presence of such widely different series of chromosome numbers
in the species of even the same genus and in genera placed under
different tribes and subtribes, indicates that the different chromosome
numbers can be derived one from the other.
Perusal of the literature (vide Table 57) reveals that with the
exception of a few genera, where the available cytological data is very
meagre, all others exhibit a wide range of chromosome numbers.
Tribe - Ocimeae
The chromosome numbers in different grougs of Lamiaceae has proved to be
of important systematic value (Harley and Heywood, 1992). Thirty three
taxa belonging to seven genera have been examined. Wide range of
chromosome numbers occur in this tribe. But in spite of these
differences, there exist a relationship between the different genera and
species as evidenced by the frequent occurrence of inter and intra
specific variations.
Ocirum L. The genus seems to be highly plastic, both numerically and
structurally. The chromosome number ranges from 2n = 22 in O, adscendens
Willd to 2n = 72 in - 0. americanum L. (vide Table 56). This wide range of
chromosome numbers may be due to the difference in number of chromosomal
biotypes belonging to the different groups (Khosla and Sobti, 1985).
Previous chromosome counts suggest that the genus tolerates a high
chromosomal diversity in it's constitution. Some stabilized diploid
numbers found are 2n = 32, 2n = 40, 2n = 48, 2n = 64 and 2n = 72 (vide
Table 57).
Among the different members of Ocimum L., O, adscendens Willd. and - 0.
americanum L. show intraspecific variations (vide Figs. 2e, 3e, 3f, 3g)
Whereas O, gratissimum L. exhibits both inter and intraspecific
variations (vide Figs. 8b, 8e, 8f, 9b, 9e, 9g) variations in chromosome
number may be due to non-disjunction, somatic reduction or even by
partial endomitosis. Non-disjunction, in the somatic tissue, involves
unequal anaphasic separation, which results in unequal distribution of
chromosomes in the daughter nuclei (Sharma, 1976).
In - 0. basilicum L. the somatic chromosome number of 2n = 48, which seem
to be constant in the four varieties studied along with the absence of
any somatic variation numbers suggest the stability of the chromosomal
biotype. The absence of somatic variation numbers might have been
probably due to the cytologically diploid behaviour and normal
disjunction of chromosomes in these polyploids, during meiosis (Riley and
Chapman, 1958).
On the contrary, - 0. tenuiflorum L.f. is characterized by interspecific
variations. The chromosome number is found to be 211x32 in the cultivars
green and purple. Whereas in the cv. purple-green and the variety hirsuta
Hook.f., it is 2n = 36 (vide Table 56). The occurrence of different
cytotypes in the sane species suggest that chromosome number in this
species is not very stable and may alter either by the duplication or
loss of individual chromosomes (Morton, 1962).
In the closely related genetical strains of - 0. gratissimum L. (2n=40) and
0. gratissirnum L. var. suavis H0ok.f. (2n = 481, there exist two - chromosomal biotypes. The existence of different chromosomal biotypes in
the same species is a remarkable feature exhibited by the genus Ocimum
L.(Pushpangadan and Sobti, 1982).
Acrocephalus Benth. The chromosome count made on the lone member
investigated, A, capitatus Benth. is 2n = 18. Cytological reports are
rare in literature, probably due to the minute chromosome size.
Orthosiphon Benth. In this genus the chromosome number 2n = 28 occur in
three taxa and 2n = 26 in two cytotypes investigated. The somatic number
2n = 24 is rare (vide Table 56). The dominance of the chromosome numbers
2n = 28 and 2n = 26 in Orthosiphon Benth. is very well established in
literature (vide Table 57).
Plectranthus L' Herit. This genus is represented by two species, one with
2n = 28 (P. - nummularius Briq.) and the other with 2n = 26 (P, wightii
Benth.) in the present study. A review of literature reveals that the
chromosome number in Plectranthus L' Herit. ranges from 2n=22 to 2n = 84,
with a high frequency for the number 2n=28 (vide Table 57).
Coleus Lour. As regards the chromosome number this genus seems to be
heterogeneous with the frequent occurrence of the somatic numbers 211~28,
2n = 32 and 2n - 48. A similar trend has also been observed in literature (vide Table 57). Somatic variations are found in most of the members of
Coleus Lour. that are also adopted asexual means of multiplication. In
vegetatively reproducing plants variation leading to speciation is caused
by chromosome alterations in the somatic tissue (Sharma, 1956; Konvicka
and Levan, 1972).
Auisochilua Wall. The chromosome counts made on the genus were 2n = 34 in
A. carnosus (L.f.) Wall and 32 in A. eriocephalus Benth., both falling in - - terms with the previous reports (vide Table 57).
Hyptis Jacq. The chromosome numbers of 2n=32 and 2n = 28 have been found
in - H. capitata Jacq. and - H. suaveolens (L.) poit. respectively. These
numbers come under the wide range of chromosome numbers (211-16 to
96)reported for the genera (Harley and Heywood, 1992). More over the most
common numbers in literature happens to be 2n = 32 followed by 28 (vide
Table 57).
Tribe Hentheae Eight taxa belonging to three genera have been examined.
The chromosome numbers of individual taxa ranges from 211-24 to 2n = 96.
This tribe is also characterized by the frequent occurrence of
intraspecific variations. The chromosome numbers and the associated
variations provides a clue in determining the status of a species
(Sharma, 1976).
Pogost- Desf. The counts made on the genus reveal the codominance of
two chromosome numbers, 2n = 64 and 2n = 32. The dominance of these two
numbers in Pogostemon Desf. is very much evident in literature (vide
Table 57). These two numbers seem to be the multiples of a common
ancestral number and hence appears to have a common origin.
EusteraUs Rafin. The chromosome number 211-30 found in - E. quadrifolia
(Benth.) Panigrahi seems to be novel. Authentic cytological reports are
not met with in this genus.
kntha L. In the present study the counts of 2n=96, 2n = 48 and 2n = 24
shows the chromosomal diversity of the genus. All the three cytotypes
were characterized by variation numbers.
Previous work on the chromosomes of Mentha L. have shown that the
species vary considerably as regards the numerical status (vide Table
57). The genus is polymorphic. Sometimes different chromosome races are
found in the same species (Darlington and Wylie, 1955). Previous records
show that the chromosome number in Mentha L. range from 2n = 18 to 2n=144
(Morton, 1956a; Harley and Brighton, 1977).
Tribe: Salvieae
S a l v i a L. It is the lone genus investigated in this tribe. Out of the
four species studied, two possess 211-22 (S. - coccinea Juss. and - S.
leucantha Cav.), one shows 2n = 16 + 0-1B ( S . - plebeia R. Br.) and the
other 2n = 44 (S, splendens Ker - Gawl). Somatic variations are common
and several chromosomal biotypes are found in a species from which
different chromosome numbers are met with. A scrutiny of the world-wide
data indicates that the chromosome number in the genus Salvia L. ranges
from 2n = 12 to 2n = 240 (Harley and Heywood, 1992). In the genus the
most frequent numbers reported were 2n = 22 followed by 2n = 14, 2n = 16,
2n = 18, 2n = 20, 2n = 32 and 2n = 44 (vide Table 57).
Tribe Larieae
Eight taxa belonging to three genera have been examined. The normal
chromosome number ranges from 2n=22 to 2n = 34. Members of this tribe are
characterized by the absence of somatic variants. Chromosome numbers
certainly appears to have played a contributory role in the evolution of
different groups in the family through speciation and morphological
diversification (Harley and Heywood, 1992).
dniso~eles R.Br. In the two members investigated in this genus, A,
indica (L.) Kuntze possess 2n - 34 and - A. malabarica R. Br., 2n = 32.
These numbers seems to have originated from a common ancestor or one from
the other. Previous studies support the dominance of these two numbers in
Anisomeles R. Br. (Vide Table 57).
Leucas R. Br. The chromosome number was found to be constant (2n=22) for
the five investigated species of the genus. This shows the stability of
the genus at the infrageneric level. 2n = 22 seems to be the predominant
number along with 2n - 28 in the literature (vide Table 57). Leonotis R. Br. The chromosome number in L, nepetifolia (L.) Ait.f. was
found to be 2n=28, which falls in agreement with the earlier reports of
2n = 24, 2n = 26 and 2n = 28 made on the genus. (Vide Table 57).
Tribe djugeae
Teucriln L. The chromosome count made on the single taxa investigated in
the tribe, - T. plectranthoides Gamble is 211~32. This seems to be a
confirmation of an earlier report made on the same taxa (vide Table 57).
b. Basic chromosome numbers in the genera studied
Basic chromosome number forms one of the widely used characters in
formulating phylogenetic speculations and hence can be considered as a
dependable and stable marker of the direction of evolution (Jones, 1970,
1974, 1978). In Angiosperms several basic chromosome numbers are involved
in various polyploid series (Grant, 1981, 1982).
A review of the world-wide data on the basic numbers of Lamiaceae reveal
that they range from X - 5 to X = 64 (Darlington and Wylie, 1955;
Fedorov, 1969; Moore, 1973). However, the most common numbers were found
to be ranging between X = 5 and X = 19 (Leshukova, 1970).
In the present investigation the basic chromosome numbers vary widely in
South Indian Lamiaceae. Out of the fifteen genera investgated three have
been found to be polybasic (Ocimum L., Orthosiphon Benth. and Coleus
Lour.), five to be dibasic (Plectranthus L' Herit., Anisochilus Wall.,
Hyptis Jacq. Salvia L. and Anisomeles R. Br.) and three genera to be
monobasic (Pogostemon Desf., Mentha L., Leucas R. Br.) In the current
study, owing to the presence of only one species, it would not be
appropriate to categorize the genera Acrocephalus Benth., Eusteralis
Rafin., Leonotis R.Br. and Teucrium L. as monobasic. Thus it appears that
the family is a highly evolved one.
A great variability in the number of chromosomes in the basic set is a
characteristic feature for the mint family (Leshukova, 1970). This
variability could possibly be the result of aneuploidy at generic level
(Fernandes and Leitao, 1984).
Both primary as well as secondary base numbers are involved in the
evolution of the fifty four taxa studied. The primary base numbers (XI)
range from 6 to 9 and secondary base numbers (X2) from 10 to 13 and 15 to
17. The basic number X2 = 12 was found in majority of the members
(24.07%) followed by the numbers X2 = 11, X1 = 7, X1 = 8 and X2 = 16 with
percentages 16.67, 14.81, 12.96 and 11.11 respectively (vide. Fig. 58).
It is likely that protopolyploidy plays an active role in the evolution
of these basic numbers (Fujita, 1970).
In Lamiaceae, the secondary base numbers might have evolved from the
primary numbers, X1 = 6-9. Grant (1981) proposes the original primary
base numbers of angiosperms to range from X = 7 to 9 and they seem to be
ancestral in the phylogenetic sense. From these primary base numbers many
polyploid series develops both by autopolyploidy and amphipolyploidy. The
latter involves both hyper and hypoploidy (Fernandes and Leitao, 1984).
Thus it can be confirmed that polyploidy had influenzed and played a role
in the evolution of base numbers, mostly of the higher order ususally by
"polyploid drop" (Darlington, 1956) and occasionally by 'polyploid lift'
(Jones, 1970).
Tribe - Ocbeae
The common base numbers of this tribe found in the present
investigation are X = 6-13, 16 and 17. Ten out of the thirty three
members investigated are based on the genome number X = 12. Other
prominent numbers are X = 7 and 8.
Ocirum L. This is a polybasic genus (Bir and Saggoo, 19851, with both
primary and secondary base numbers observed in the present investigation.
A great diversity (X = 8-12) is observed in this regard, with half of the
investigated taxa being based on X = 12. These base numbers might have
evolved from the primitive base number X = 6, through dysploidy and
polyploidy at different levels (Sobti and Pushpangadan, 1977). Literature
evidences reveal the dominance of two common base numbers (X = 8 and
X = 12) in Ocimum L. (Sobti and pushpangadan, 1977; Singh and Sharma,
1982; Khosla, 1989).
Ocimum adscendens Willd. possess the secondary basic chromosome
number, X2 - 11. This might have been formed by amphipolyploidy (Grant,
1981) from the primary base numbers X1 = 5 and 6. The chance for the
evolution through ascending or descending dysploidy (Love, et al., 1957)
from X2 = 10 and X2 = 12 cannot be ruled out. - 0. americanum L. (2n=72)
exhibits secondary major protopolyploidy during the course of evolution
of it's base number X2 = 12 from the corresponding primary number (XI
= 6). Monobasic status (X2 = 12) was evident in 0. basilicum L., with all - the four varieties exhibiting doubling of the primary basic set X1 = 6.
The dibasic nature of - 0. gratissimum L. (2n = 40) with X2 = 10 and - 0.
gratissimum L. var. suavis H0ok.f. (2n = 48) with X2 = 12 might have been
due to independant autopolyploidy (Stebbins, 1971) from primary base
numbers X1 = 5 and 6 respectively.
The different cultivars and a variety of 0. tenuiflorum L.f. are set upon -
two primary base numbers. The basic number X1 = 8 was found in 0. -
tenuiflorum L.f. cv. green and cv. purple ( 2 ~ 3 2 each). Whereas in 0. - -
tenuiflorum L.f. cv. purple -green and 0. tenuiflorum L.f. var. hirsuta -
Hook.f., the basic set was found to be X1 = 9. To establish an ancestral
count among these two primary base numbers is a matter of confusion. The
primary base number X1 = 9 might have originated from X1 = 8 either by
progressive evolution or the vice versa by primary reduction (Jones,
1970).
Acrocephalus Benth. The basic chromosome number found in the lone member
investigated, A, capitatus Benth. was primary in nature with X1 = 9
chromosomes.
Orthosiphon Benth. This genus reveal the existence of three base numbers
out of the six members investigated. They are X1=6 in - 0. thymiflorus
Roth. (2n=24), X1=7 in - 0 . grandiflorus Bold cv. lilac, cv. white and 0,
pallidus Royle (each with 2n=28) and X2-13 in O, glabratus Benth. and - 0.
glabratus var. ~arviflorus Benth. (each with 2n=26). Morton (1962)
considers X1 = 7 as the original primary basic number of this genus. Then
the origin of X1 = 6 can only be through a diminution of the base number
(Stebbins, 1950). The secondary basic number X2=13 found in - 0. glabratus
var . parviflorus Benth. might be a case, evolved through polyploidy
followed by aneuploidy or dibasic polyploidy (Gill, 1970) involving the
primary base numbers X1 6 and 7.
Plectranthus L'Herit. In the present study this genus seems to be dibasic
with the primary base number X1 = 6 in P, wightii Benth. (2n=24) and X1 =
7 in - P. nummularius Briq. (2n=28). The common base number of Plectranthus
L'Herit is X = 6 (Darlington and Wylie, 1955). However deWet (1958b) is
of the opinion that X = 7 also exists. In Plectranthus L'Herit., the
primary basic number X1 = 7 can be the model number from which X1 = 6 is
derived by reduction (Morton, 1962).
Coleus Lour. The genus is polybasic (Bir and Saggoo, 1982a) with both
primary as well as secondary base numbers existing in it's members. The
primary basic number XI = 7 was found in - C. forskohlii Briq. as well as
C. zeylanicus (Benth.) Crarner and X1 = 8 in C. aromaticus Benth. and C. - - -
aromaticus Benth. var. variegata ( 2 ~ 3 2 each). The secondary base number
(X2 = 12) was found in - C. blumei Benth. var. verschaffeltii, - C.
rehneltianus Berger (211x48 each) and in - C. parviflorus Benth. (2n=72).
The primary basic numbers XI = 6, X1 = 7 and X1 = 8 have long been known
in Coleus Lour. (Darlington and Wylie, 1955). The ancestry of these
primary base numbers is a matter of considerable debate. Here X = 6 may
be considered as the primitive model base number from which XI = 7 and X1
= 8 might have been arisen, since an increase in the basic chromosome
number is easier to envisage and is likely to cause less unbalance than a
decrease (Darlington, 1963). The secondary basic number X2 = 12 has been
well established in Coleus Lour. (Bir and Saggoo, 1979, 1982a). This
higher basic number is considered to be derived f r o m lower numbers by
autopolyploidy (Stebbins, 1966).
BnisochFZue Wall. The genus exhibit two secondary basic numbers, one
with X2 = 16 in A, eriocephalus Benth (2n = 32) and the other with X2 =
17 in - A. carnosus (L.f.1 Wall. (2n=34). The secondary basic chromosome
number of X2 = 16 might have derived it's origin by autopolyploidy
(Fernandes and Leitao, 1984) from the primary base figure of X1 = 8.
Where as X2 = 17 might have formed by amphipolyploidy (Grant, 1981) from
the primary numbers XI = 8 and X1 = 9 or either through a 'polyploid
drop' (Darlington, 1956) from the primary base number X1 = 9 or through
a 'polyploid lift' (Jones, 1970) from X1 = 8.
Hyptis Jacq. The basic numbers found in the genus are primary in nature
with X1 = 7 and X1 = 8 and hence the genus appears to be dibasic. Morton
(1962) established these two numbers as the primary basic figures of
Hyptis Jacq. These numbers might have originated one from the other
through an increase (Darlington, 1963) or through diminution (Stebbins,
1950) of primary base figures.
Tribe Hentheae
In this tribe, the common base numbers found are 12, 15 and 16, all of
them being secondary in origin. Auto and amphipolyploidy seem to be
involved in the evolution of various members of this tribe.
Pogostemon Desf. The genus is monobasic with the secondary number X2 =
16, found in all the four investigated members. The existence of these
basic numbers were confirmed by Bir and Saggoo (1982a). This base number
might have been formed through autopolyploidy (Mehra and Bawa, 1969)
during evolution.
Eusteralis Rafin. The basic number found in the lone member investigated,
E. quadrifolia (~enth.) Panigrahi (2n=30) is X2 = 15. This secondary -
basic number is formed either through polyploidy followed by aneuploidy
or dibasic polyploidy (Gill, 1970) involving the primary base numbers
X1 = 7 and X1 = 8.
tkntba L. All the three investigated species are multiples of the
secondary basic chromosome number X2 = 12. Evidences from the literature
(Ikeda and Udo, 1963; Harley and Brighton, 1977) reveal the predominance
of this base number in the genus. This secondary base number can be
formed only through autopolyploidy (Stebbins, 1966) from the primary
basic number, X1 = 6.
Tribe - Salvieae
The basic chromosome numbers found in the investigated members of this
tribe include X1 = 7, X1 = 8, X2 - 11, X2 = 16 and X2 = 17. Both
ascending and descending dysploidy, auto polyploidy and amphipolyploidy
were found to be responsible for the evolution of various taxa coming
under this tribe.
S a l v i a L. The genus appears to be dibasic with both primary as well as
secondary base numbers found in the present investigation. When the
primary number X1 = 8 was found exclusively in S. plebeia R. Br., the - secondary number X2 = 11 was prevalent among S, coccinea Juss., S. -
leucantha Cav. and - S. splendens Ker-Gawl.
The genus Salvia L. is polybasic (Palomino, et al. 1986). Scrutiny of the
world-wide data indicates that the basic numbers range from X = 6 to 19
(Saggoo and Bir, 1986). However, the numbers 6, 7, and 8 are considered
as primary base numbers (Gill 1971 a). According to Afzal - Rafii (1976)
the base numbers prevalent in Salvia are X1 = 7 and X1 = 8, the latter
might have originated from the former as a result of progressive
evolution (Afzal-Rafii, 1971). The primary base numbers X1 = 5 and X1 = 6
seems to be involved in the amphipolyploid origin (Grant, 1981) of the
secondary basic chromosome number X2 = 11. It can also be formed by
ascending or descending dysploidy (Love, et al., 1957).
Tribe - Lamieae
Both primary as well as secondary base numbers seem to be prevalent in
this tribe. Dibasic polyploidy and descending or ascending dysploidy are
involved in the evolution of secondary base numbers.
Anisomeles R. Br. This genus exhibits the secondary basic number X2 = 16
in A, malabarica R. Br. (2~32) and X2 = 17 in - - A. indica (L.) Kuntze
(2~34). The former number arises by autopolyploidy (Stebbins, 1950) of
the primary base number X1 = 8 and the latter by dibasic polyploidy
(Gill, 1970) of the primary base numbers X1 = 8 and X1 = 9. A chance for
the origin of X2 = 17 through a 'polyploid lift' (Jones, 1970) of the
primary number X1 = 8 or even by a 'polyploid drop' (Darlington, 1956)
from XI = 9 cannot be ignored.
Leucas R. Br. All the five cytotypes of the genus possess the same
secondary base number X2 = 11. Literature surveys (Vij and Kashyap,
1976b; Bir and Saggoo, 1979, 1982a; Saggoo and Bir, 1986) confirms the
existence of X2 - 11 in Leucas R. Br. Chance for the occurrence of
amphipolyploidy (Grant, 1981) from the primary base numbers X1 = 5
and X1 = 6 as well as ascending or descending dysploidy (Love, et al.,
1957) from the respective secondary base numbers are equal during the
process of evolution.
Leonotis R. Br. The base number found in L. nepetifolia (L.) Ait.f. is
X1 = 7, which is primary in nature and well supported by an earlier study
(Morton, 1962).
159
Tribe Ajugoideae
Teucrium L. - T. plectranthoides Gamble (2n=32), the lone member
cytologically screened in this tribe reveals the presence of the primary
basic chromosome number X1 = 8, which has been confirmed earlier by
Koudysheva and Akhmed - Zade (1968).
c. Polyploidy in the genera investigated
Polyploidy is a very widespread cytogenetic phenomenon found in over 30%
of dicotyledons and 50% of monocotyledons (Love and Love, 1975). It is a
mechanism which involves multiplication of the whole chromosome
complement and there by an increase of gene number and variety, producing
radically different well adopted genomes (Stebbins, 1950; Gottschalk,
1979).
Polyploids are considered to be more hardy and adaptable to extreme
climatic conditions (Hagerup, 1931; Tischler, 1935; Love and Love, 1942b,
1943). However the theory of hardiness was criticized by Bowden (1940),
Clausen, et al., (1940, 1945) and Nielsen (1947).
Polyploidy is one of the best genetical and evolutionary processes, which
has greatly contributed to speciation and evolution of higher plants
(Gottschalk, 1985). This is mainly due to the ability of polyploids to
increase the chances of fertilization by breaking reproductive barriers,
which permits natural selection and establishment of species even under
adverse environmental conditions (Winge, 1917; Stebbins, 1971, 1974).
There can be no doubt that polyploid species are highly successful. A
comparison of the geographic distribution of polyploids and diploids in
plants show a greater adaptability of polyploids (Love and Love, 1943).
However the evolutionary potentialities of a diploid are likely to be
greater, in the long run (White, 1937).
The family Lamiaceae is characterized by a relatively high frequency of
polyploidy (Morton, 1962; Fernandes and Leitao, 1984). Studies conducted
on fifty four taxa reveals the dominance of polyploids (62.96%) over
diploids in South Indian Lamiaceae. Out of these, 57.41% are tetraploids
followed by hexaploids (3.7%) and octaploids (1.85%) (vide Fig. 59).
The present study reveal that herbaceous elements predominate as compared
with the shrubby elements of the family. Growth habit is one of the
factors which influence the frequency of polyploids in
angiosperms(Baquar, 1976). Correlating polyploidy with life form and
growth habit, it is clear that the highest frequency of polyploids occur
in perennial herbs (79.41%), lowest in annual herbs and undershrubs
(2.94% each) and intermediate in perennial undershrubs (14.71%) vide Fig.
60). In Angiosperms, the highest percentage of polyploidy was found in
perennial herbs while annuals and woody plants have lower percentage of
ploidy (Stebbins, 1938; Fagerlind, 1944) The origin of perennial habit,
as well as accessory methods of vegetative reproduction is considered to
be a direct consequence of polyploidy (Muntzing, 1936; Gustafsson, 1947a,
1948).
The various genera which show a predominance of polyploidy include Ocimum
L., Orthosiphon Benth., Coleus Lour., Hyptis Jacq., Pogostemon Desf.,
Mentha L., Salvia L., Leonotis R.Rr. and Teucrium L.
Tribe - Ocimeae
Different degrees of polyploidy has been observed in this tribe. Out of
the thirty three taxa investigated, twenty seven (81.82%) exhibit
polyploidy, with the majority being tetraploids, found in 25 members. The
role of polyploidy in the mechanism of speciation is obvious in the tribe
Ocimeae (Bhattacharya, 1978b).
Ocimum L. The cytological datas obtained from the genus Ocimum L.
indicated that eleven out of twelve members (91.67%) are polyploids.
Ocimum adscendens Willd. (2n = 22), the lone diploid representative,
exhibits a tetrasomic variant number of 24 (Fig. 2e) This might have been
probably caused by multipolarity that leads to unbalanced chromosome
complements (Brachet, 1957). At the molecular level the predisposition of
polyploidy shown by diploid individuals may be due to a change in the
gene expression and enzyme behaviour which causes a disturbed metabolic
regulation and consequently the abnormality of development. (Albuzzio, et
al., 1978).
Among the polyploid members, tetraploidy is found in all except - 0.
americanum L. (2n = 721, where the ploidy level is 6x. This hexaploid
might have been derived from a polyploid series based on the 'basic set of
12 chromosomes through autopolyploidy (Fernandes and Leitao, 1984).
All the varieties of - 0. basilicum are characterized by the absence of
variants. The stability of the chromosome number ( 2 ~ 4 8 ) indicates that
distinctly well differentiated genomes are involved in the origin of
these polyploids (Koul and Gohil, 1991).
162
0 . gratissimum L. (211-40) and 0. gratissimum L. var suavis Hook. f.(2n = - --
48) exhibit aneuploid somatic variants. Aneuploidy is very common in the
family at both generic and specific levels (Morton, 1962). It is caused
by the gain or loss of one or more chromosomes from the haploid set
(White, 1937).
The cultivars of - 0 . tenuiflorum L.f. exhibit interspecific polyploidy.
This is due to the existence of two different sets of chromosome numbers
2n = 32 and 36 derived through two different series, from the primary
base figures X1 = 8 and X1 = 9, by autopolyploidy (Clausen, et al.,
1945).
A perusal of the literature shows that so far all the investigated =urn
species are at polyploid level. This shows that polyploidy has played a
major role in the speciation of this genus (Morton, 1962; Pushpangadan,
et al., 1975).
Orthosiphon Benth. It is a genus, which was found to exhibit tetraploidy
in four out of the six members (66.67%) investigated. Here the cause for
polyploidy in the two cultivars of - 0 . grandiflorus Bold (2n = 28 each) - 0.
pallidus Royle (2n = 28) - 0. thymiflorus Roth. (2n = 24) may be due to the
parallel evolution of two basic figures X1 = 7 and X1 = 6 and formation
of multiples which develop into two series of polyploids during the
course of evolution (Fernandes and Leitao, 1984).
Plectranthus L' Herit. The two members investigated in the genus, namely
P. nummularius Briq. (211~28) and P. wightii Benth. (2n = 24) are - --
tetraploids. The genus shows predominance of polyploidy (deWet, 1958b).
163
Coleus Lour. The various cytotypes of the genus Coleus Lour. exhibits
polyploidy and varying degrees of somatic variations. The delicately
balanced system of gene interaction in tetraploids is disturbed by the
doubling of chromosomes, resulting in the formation of somatic variants
(Kuckuck and Levan, 1951).
C. aromaticus Benth. and C, aromaticus Benth. var. variegata are - - tetraploids with a basic set of X1 = 8 chromosomes. In both these plants
many somatic variants are present, which are multiples of the lowest one.
When such a phenomenon exists with in a single species of plant, it is
possible that they have arisen by autopolyploidy (White, 1937). The other
variant numbers can be hyperploids (Khush, 1973).
Members like - C. forskohlii Briq. and - C. zeylanicus (Benth.) Cramer
(2n=28 each) are autotetraploids based on the base figure X1 = 7. The
somatic variation numbers of - C. forskohlii Briq. might have been formed
through ascending or descending dysploidy (Love, et al. 1957). - C.
zeylanicus (~enth.), Cramer exhibits both aneuploid variant numbers (2n =
30 and 2n = 32) as well as multiples of the basic set of 7 chromosomes
(2n = 14 and 2n = 56). This shows the role of aneuploidy and
autopolyploidy that have played an important role in the evolution of the
taxa (Ramachandran, 1967).
The tetraploid taxa based on the somatic chromosome number 2 ~ 4 8 ,
blumei Benth. var. verschaffeltii, - C.laciniatus Benth. and C,
rehneltianus Berger may probably have an allopolyploid origin (Hakeem and
Rife, 1966). These three taxa also exhibit dysploid somatic variation
numbers. Dysploidy is quite a common feature found in Lamiaceae (Marrero,
1992).
The vegetatively propagated taxa, - C. parviflorus Benth. exhibits a high
degree of polyploidy (6x), probably due to the failure of cell division
in the first instance (White, 1937). It also exhibits varying degrees of
autopolyploid and aneuploid variants, a common feature of this family
(Fernandes and Leitao, 1984).
Byptis Jacq. The obnoxious weed - H. capitata Jacq. (2n=32) and it's ally
H. suaveolens (L.) Poit. (2n = 28) are tetraploids. There are several -
reports on records, where weedness is often associated with polyploidy
(Cain, 1944).
Tribe llentheae
This tribe was found to attain very high degrees of polyploidy in some
members and four out of eight members exhibit polyploidization. Most of
these polyploids are resorted to vegetative means of reproduction, an
adaptive feature exhibited by polyploids (Fagerlind, 1944; Stebbins,
1947).
Pogostamon Desf. Out of the four taxa investigated in this genus, P,
benghalensis (Burm.f.) Kuntze and P- heyneanus Benth. are tetraploids
with 2 ~ 6 4 . The diploids, P, purpurascens Dalz. and - P. vestitus Benth.
possess a somatic number of 2n = 32 each. Eventhough both the polyploid
and diploid taxa of Pogostemon Desf. are multiples of the basic figure X
= 16, the higher levels of ploidy attained by - P. benghalensis (Burm.f.)
Kuntze and & heyneanus Benth. probably point towards their greater
adaptability. The aneuploid and hypoploid variations found in these
polyploid taxa reveal the ongoing genetic and evolutionary processes
which may help to break the reproductive barriers enabling natural
selection and thereby speciation (Stebbins, 1974).
Hentha L. various levels of ploidy have been observed in the three
cytotypes investigated in this genus. Out of these - M. arvensis L. is an
octaploid, exhibitingfiypo+loid somatic variation numbers.
The octaploid status of & arvensis L. was confirmed earlier by Ikeda and
Seiroku (1966) and Tyagi (1986). Aneuploid numbers in & arvensis L. has
been reported earlier by Sharma and Bhattacharya (1959a), Ouweneel
(1968) as well as Tyagi and Naqvi (1987).
M. rotundifolia (L.) Huds. is characterized by it's diploid status and - occurrence of aneusomatic variation numbers. These numbers are multiples
of the lowest complement there by exhibiting proto-autopolyploidy (White,
1937). Polyploidy, aneuploidy and aneusomaty were found to be effective
in the evolution of various species of Mentha L. (Kundu and Sharma,
1985).
The cytotype - M. spicata (L.) Huds. exhibits tetraploidy (2n=48) as well
as aneuploid, hypoloid and hyperploid variations. Aneuploidy has been
reported on & spicata L. Huds. by Ruttle (1931a) and Nagao (194la).
Moreover the vegetative means of reproduction, which is prevalent in the
investigated taxa seems to bear a correlation with the high degrees of
polyploidy. Polyploidization might have led to the establishment of new
gene combinations that have triggered off new developmental changes
leading to a later shift towards the a sexual mode of reproduction
(Stebbins, 1980; Gustafsson, 1947b). Thus polyploidy and aneuploidy was
found to play an important role in the evolution of Mentha L. (Makarov
and Reznikova, 1972; Ouweneel, 1968; Pogan. et al., 1986).
Tribe - Salvieae
In contrast to the other tribes, it shows a lower percentage of
polyploids among the members studied.
Salvia L. It is a genus of much interest from the cytological point of
view. Eventhough polyploidy is prevalent only in one out of the four
species studied, variants occur in all the species, exhibiting natural
aneuploids from nullisomy to hexasomy. S, splendens Ker-Gawl. was the
only member to show tetraploidy. This was confirmed by earlier findings
(Bhattacharya, 1973; Bir and Saggoo, 1982a, 1985). Dysploidy was also
found in the different members of the genus (Afzal - Rafii, 1972, 1976).
The known trends of evolutionary development in the genus are dysploidy
(descending and ascending) with in the range of the genus and with in the
range of a single taxon, polyploidy and aneuploidy (Markova and Ivanova,
1982a; Palomino, et al., 1986)
Leonotis R.Br. The lone taxa investigated, - L. neptetifolia (L.) Ait. f.
shows tetraploidy from the base figure X1 = 7. Thus it seems to be a case
of autopolyploidy (Clausen, et al., 1945).
Tribe Ajugeae
Teucrium L. The single taxa investigated in the tribe, plectranthoides
Gamble, reveal the existence of autopolyploidy from the basic figure X1 =
8. An increase in the number of chromosomes through autopolyploidy
provides increased possibilities for new gene combinations, which are of
considerable importance in evolution (Reese 1961, 1966).
d) Karyowrphometrical analysis
In Angiosperms, the species of several families, both dicotyledons and
monocotyledons are found to exhibit a direct relationship between their
phylogeny and the chromosome constitution. The chromosomes being the
carriers of heredity, both structural and numerical changes in them can
influence the genetic-evolutionary process at work than do any other type
of changes. A detailed information regarding the chromosome architecture
in higher plants can thus serve as a useful tool to understand their
systematic relationships and for tracing the trend and direction of their
evolution (Love and Love, 1975).
Some of the major karyotype characteristics of considerable evolutionary
and taxonomic significance are (1) differences in the absolute chromosome
size, (2) differences in the position of the centromere, (3) differences
in total chromatin length (4) differences in karyotype formula and (5)
number as well as position of satellites.
The general feature noted in the family Lamiaceae is the wide range of
chromosomes with graded symmetrical karyotypes (vide Fig. 56), as has
been found in an earlier study. (Kundu and Sharma, 1988 a). However the
chromosome complements in South Indian Lamiaceae members differ in minute
karytypic details (vide Table 56). With regards to the gross morphology,
chromosomes are nearly submetacentric to nearly metacentric in nature.
The chromosomes range from 4.2 p to 1.0 p in length. The chromosomes
with secondary constriction ranges from two to six in number. The average
chromosome length (ACL) varies from 2.74 ,u to 1.12 p. The total chromatin
length (TCL) shows a very wide variation with 24.4 ,u being the minimum
and 154.8 p being the maximum value. The disparity index (DI) values
range between 22.22 and 50. The coefficient of variation ranges from
12.53 to 30.00 and total centromeric index (TFX) ranges from 35.83 to
45.07.
These variations found in the karyotypic parameters suggest that South
Indian Lamiaceae members are characterized by symmetrical to slightly
asymmetrical karyotypes. An increase in the range of chromosome length as
well as the increase of submetacentrics at the expense of metacentrics is
accompanied by an increase in the coefficient of variation leading to
asymmetry (Stebbins, 1958). Thus karyomorphological studies are of
considerable importance in order to throw light on the phylogenetic
relationship among taxa of flowering plants (Iwatsubo and Naruhashi,
1991).
Tribe - Ocimeae
Ocirum L. This major genus is characterized by a general homogeneity
observed in the minute details of the karyotype (Singh and Sharma, 1982).
The range of chromosome length (RCL) and their general morphology is the
same as has been noted for the family.
A remarkable feature found in all polyploid taxa of Ocimum L. is the
presence of four chromosomes with satellite or secondary constrictions
inspite of the difference in chromosome number. Thus it is clear that
numerical variations do not involve the satellited chromosomes. However
the micromorphology of these chromosomes varies from cytotype to
cytotype, indicating that structural changes have been operating in their
evolution. In spite of the evidence of the so called minor structural
variations, the presence of four nucleolar chromosomes in all the
polyploid forms of Ocimum L. indicate that these nucleolar chromosomes
may contain vital genes for the survival of the genus, which are
essential and not affected by the structural changes noted so far (Singh
and Sharma, 1981a).
The differences observed in the karyotype formula (KF), average
chromosome length (ACL) and chromatin length of basic complement (CLBC)
among different taxa might have been probably due to structural
alterations of chromosomes (vide Table 56; Fig. 61). Speciation depends
more on chromosomal rearrangements and mutation of individual genes than
on changes in the total amount of genetic content (Stebbins, 1959).
The high values of average chromosome length and chromatin length of
basic complement shown by - 0. adscendens Willd. and 0, americanum L. (vide
Table 56) probably show their primitiveness. Whereas, the lower values
for these parameters found in O,tenuiflorum L.f. cv. purple and CV.
purple-green denotes their highly evolved nature. A decrease in chromatin
length is one of the factors responsible for evolution of higher plants.
(Babcock and Cameron, 1934).
The high disparity index (DI) value found in - 0. americanum L. followed by
0. basilicurn L. var. purpurascens Benth. and var. pilosum Benth. - correspond to the heterogeneous assemblage of chromosomes in these taxa
(vide Fig. 56).Where as, the lower values of DI found in other members
point towards the general homogeneity found in various species of Ocimum
L. Normally a low disparity index value corresponds to the homogeneity of
chromosomes in most of the higher as well as lower plants (Mohanty, et
al., 1991).
In addition to the above, the high mean centromeric index (TF%) value
(vide Table 56) undoubtedly confirms the primitive status of - 0.
americanum L. A high TF% value represent a highly symmetric karyotype,
which is a primitive condition (Stebbins, 1959). However the
comparatively higher variation coefficient and lower TF% values exhibited
by - 0. tenuiflorum L.f. cv. purple shows that it represent the climax of
evolution among the different taxa investigated in Ocimum L.
In - 0. basillcum L., the karyotype of the variety purpurascens Benth.
possess a high number of medium-sized chromosomes (Type C) and low number
of small-sized chromosomes (Type Dl. Whereas, it is just the reverse in
the variety glabratum Benth. (vide Fig.56). A reduction of chromatin
length in - 0. basilicum L. var. glabratum Benth. appeared to have taken
place due to chromosomal rearrangement and the role of heterochromatic
segments during the process of evolution. Here the comparatively inert
heterochromatic segments might have been deleted resulting in the
shortening of the chromosome size (Darlington and Mather, 1950).
Moreover, the high values of average chromosome length, total chromatin
length, chromatin length of basic complement and TF% found in - 0.
basilicum L. var. purpurascens Benth. and the corresponding low values
found in - 0. basilicum L. var. glabratum Benth. further confirms that both
these are at two extremes in the evolutionary pathway (vide Table 56).
The decrease in chromatin length and the abundance of submetacentric
chromosomes with in a single complement are advanced karyomorphological
features (Babcock and Cameron 1934).
Concrete conclusions cannot be drawn from the karyomorphological features
exhibited by the closely related genetic strains of 0. gratissimum L. -
(2n=40) and - 0. gratissimum L. var. suavis H0ok.f. (2n=48). SQ it seems
likely that they represent two different lines as has been found in the
evolution of their basic numbers.
At the infraspecific level, O, tenuiflorum L.f. cv. purple and cv. purple
green are characterized by an increase in number of small sized
chromosomes (Type D) and the low values for the major karyomorphometrical
parameters except variation coefficient (vide Fig. 56, 61; Table 56).
Thus these observations are in advocacy with their highly evolved status.
The phylogenetic changes occurring in the length of the chromosomes oculd
cause shortening in size of one of the arms leading to the shifting of
the centromeric position as well as reduction in the absolute length of
the chromosome (Delauney, 1926).
Acrocephalus Benth.
In this genus, the low disparity index (DI) value reveals the homogeneity
of the chromosome complement. The high T F % found in conjunction with a
low variation coefficient value (vide Fig. 62; Table 5 6 ) reveal that
A. capitatus Benth. is a primitive plant. A high T F X and low variation - coefficient values correspond to the primitive status in the evolution of
flowering plants (Stebbins, 1971).
Orthosiphon Benth.
The comparatively high values for range of chromosome length ,average
chromosome length, total chromatin length and TF% found along with a low
variation coefficient (vide Fig. 62; Table 56), which occur in the
karyotype of - 0. glabratus Benth. and 0. thymiflorus Roth. point towards -
primitiveness. In addition to these, the small sized chromosomes (Type D)
are found to be absent in their karyotype. This along with a low
disparity index value in them show their homogeneity. Long metacentric
nearly equal chromosomes with a high mean centromeric index value were
found in comparatively primitive taxa (Levitsky, 1931). However, in - 0.
pallidus Royle,the lower values for range of chromosome length, average
chromosome length and chromatin length of basic complement found in
association with a high variation co-efficient and comparatively low TF%
value point towards an evolutionary advance.
At the infraspecific level, a numerical increase in small sized (Type D)
chromosomes and comparatively low values of average chromosome length and
total chromatin length (vide Fig. 56, 62; Table 56) found in Lglabratus
var. parviflorus Benth. and - 0. grandiflorus - Bold. cv. - lilac show their
evolved nature than their allys - 0. glabratus Benth. and - 0. grandiflorus
Bold cv. - white. This has been further confirmed by the higher variation
coefficient value found in O, glabratus var. parviflorus Benth. and a low
TF% value found in - 0 . grandiflorus Bold. cv. lilac. Thus these minute
structural alterations might have been responsible for evolution. The
morphological evolution of chromosomes involves a change from uniform
length of chromosomes to differences, change of median centromeres to
submedian centromeres and reduction in chromosome length (Jones, 1978).
In addition to these, differences are also found in the number of SAT-
chromosomes (vide Fig.56; Table 56). The secondary constriction
constitute an important feature of chromosome in karyomorphological
studies at the infraspecific level (~ewis and John, 1963). The presence
of only one pair of SAT - chromosomes in the tetraploid taxa, - 0.
grandiflorus Bold cv. lilac and cv. white may be due to the loss of one
pair of chromosomes during the course of evolution. It also seems that
the amount of nucleolar genes contained with in the satellited
chromosomes is sufficient to meet the need of the ribonucleoprotein
metabolism as has been found in other plants (Singh and Sharma, 1983).
Plectranthus L'Herit. The lower values for range of chromosome length,
average chromosome length, total chromatin length as well as chromatin
length of basic complement and a comparatively high value of variation
coefficient (vide Fig. 62; Table 56) found in P, nummularius Briq., than
it's relative - P. wightii Benth. probably indicate an evolutionary
advance. Moreover, an increase in chromosome number is observed in 11,
nummularius Briq. (vide Fig. 56). Reduction in chromosome size is
apparently a consequence of polyploidy, since it is an adaptation to a
decrease in size of the cell or to an increase in number of chromosomes
(Darlington, 1956).
Coleus Lour. The different species of Coleus Lour. are exceedingly
variable both in chromosome numbers and in morphometric characters of
their karyotypes. This variability of the genus evidently reflects an
important side of it's evolution, apart from it's morphological
differentiation, on which classification is based.
In C, aromaticus Benth. the higher number of nearly median centromeres
and the higher values for all the other karyomorphological parameters
(vide Fig. 56, 63; Table 56) reveal it's primitiveness. In contrary, the
karyotypic data found in - C. aromaticus Benth. var. variegata speaks off
it's evolutionary advance.
On comparing the karyomorphology of cytotypes with 2n = 28 chromosomes,
C. zeylanicus (Benth.) Cramer exhibits a higher number of short sized - (Type D)chromosomes, higher value for variation co-efficient (V.C.) and
lower values for other karyomorphological data (vide Fig. 56, 63; Table
56) Where as, in - C. forskohlii Briq. it is just the reverse. The advanced
nature of - C. zeylanicus (Benth.) Cramer and the primitive status of - C.
forskohlii. Briq. is very much evident. Similarly - C. laciniatus Benth.
pssesses a higher number of short sized (Type D) chromosomes and lower
values for other karyotypic details (vide Fig. 56, 63; Table 56) when
compared with other members based on 2 ~ 4 8 . Hence - C. zeylanicus (Benth.)
Cramer seems to be evolved. The structural rearrangements related to
favourable position effect and gene linkage, which are likely to confer
adaptive advantage, may increase the asymmetry of karyotype (Stebbins,
1958).
C. parviflorus Benth., a hexaploid is characterized by the presence of 6 -
chromosomes with secondary constrictions (vide Fig. 56). It has been very
well established that each basic set of chromosomes has at least one
chromosome with secondary constriction (de Mol, 1926b; Bhaduri, 1944).
Moreover the lesser range of chromosome length and low variation
coefficient value and the comparatively higher TF% value indicate their
probable primitiveness (vide Table 56; Fig. 63).
Thus in the genus Coleus Lour., structural alterations seem to have a
major role during the course of evolutiqn and these changes might have
been probably due to pericentric inversion and unequal translocation in
the chromosomes (Stebbins, 1971).
175
dnisochilus Wall. Karyotypic analysis conducted on the two diploid
species of this genus reveal the homogeneity, as was evidenced by their
almost similar karyotype formula (KF) and comparatively higher value of
disparity index (DI). However, differences are noted for average
chromosome lenth, total chromatin length, chromatin length of basic
complement, variation co-efficient and TF%. (vide Fig. 64, Table 56). The
higher value among these being shown by A, carnosus (L.f.) Wall., thereby
exhibiting it's primitiveness. Karyotype with lesser range of chromosomal
length is found in comparatively primitive genera (Levitsky, 1931).
Hyptis L. Karyomorphometrical surveys on the genus are rare, probably due
to the small chromosome size. Both the tetraploid taxa investigated are
characterized by homogeneous graded karyotypes. (vide Fig. 56). In - H.
capitata Jacq. all the karyometrical values seem to be comparatively
higher than that in - H. suaveolens (L.) Poit. (vide Fig. 64; Table 56).
Such variations are important from the evolutionary viewpoint. Thus
primitiveness is more pronounced in & capitata Jacq. than suaveolens
(L.) Poit. This is probably due to karyotypic rearrangements which is an
initial step in speciation as has been found in other plants (Clausen,
1951).
Pogost-n Desf.
With in the genus Pogostemon Desf., the tetraploid taxa showed more
similarities among them than differences when compared with the diploid
taxa. The polyploid plants, P, benghalensis (Burm.f.) Kuntze and - P.
heyneanus Benth. showed homogeneous symmetrical karyotypes, a primitive
condition. However, the lesser values of average chromosome length,
chromatin length of basic complement, disparity index and TF% (vide Table
56., Fig. 64) advocates their evolved nature. Among the karyotypes of
the diploid forms, - P. vestitus Benth. exhibited a higher number of short
sized (Type D) chromosomes, comparatively low values of average
chromosome length, total chromatin length as well as TF% along with a
high value of variation coefficient. Therefore - P. vestitus Benth. can be
considered as more advanced than - P. purpurascens Dalz. Structural
alterations of chromosomes have a major role towards speciation (Singh
and Sharma, 1982) .
A remarkable feature noted in the four taxa of Pogostemon Desf. is the
presence of 4 SAT-chromosomes (vide Fig. 56), irrespective of their
ploidy level. Thus it has become undoubtedly clear that in some plants
the number of nucleolar chromosomes maynot increase proportionately with
the concomittant increase in ploidy (Singh and Sharma, 1983).
Eusteralis Rafin.
The karyotype of - E. quadrifolia (Benth.) Panigrahi seems to be formed of
very small chromosomes, symmetrical and with a very low range of
chromosome length (vide Fig. 56). Alongwith these primitive features, it
exhibits some advanced evolutionary features like low values for average
chromosome length, chromosome length, chromatin length of basic
complement and TF% (vide Fig. 65; Table 56). So the evolutionary status
of this taxa appears to be enigmatic.
Mentha L.
In the diploid, tetraploid and octaploid species of Mentha L., the
karyotype seems to be symmetrical and homogeneous (vide Fig. 56). The
presence of only two pairs of chromosomes with secondary constrictions in
a11 the species probably show that two pairs of nucleolar chromosomes are
essential for the existence and stability of the different species of
Mentha L. In some plants, SAT - chromosomes are not affected by numerical variations (Kundu and Sharma, 1985). However,cryptic structural changes
have been operating in their evolution. Among the three taxa, & arvensis
L. was found to exhibit a comparatively low TF% and high variation co-
efficient value (vide Fig. 65, Table 56). This correspond with the
evolutionary advancement of this plant. Karyotypic details also points
out the evolved status of - M. spicata (L . ) Huds. However, the diploid taxa
M. rotundifolia (L.) Huds. with a low average chromosome length, and low -
disparity index value and a high TF% seem to be rather primitive in the
evolutionary history of the genus. The centromeric index is a karyotype
characteristic independant from chromosome number. A low mean centromeric
index value suggest a highly advanced karyotype and a high value
represents a primitive karyotype (Vasil'eva, et al., 1985).
Salvia L. The genus exhibit differences as regards chromosome
morphology. In the diploid taxa based on 2n = 22, S. coccinea Juss. shows - -- comparatively lesser values for range of chromosome length, average
chromosome length, total chromatin length, chromosome length of basic
complement and TF.% (vide Fig. 65, Table 56) than S. leucantha Cav. - Eventhough a low range of chromosome length is a primitive character,
all the other karyomorphological data, along with a high variation
coefficient value reveal it's advanced status.
The karyotype of S. plebeia R.Br. is characterized by a higher variation - coefficient and a lower value for chromatin length of basic complement.
These two parameters are sufficient to place it among the advanced
members of Salvia. L. In the present study, B-chromosome are detected
only in the chromosome complement of Salvia plebeia R. Br., (vide Fig. 45
b) ranging from 0-1 in number. This B-chromosome might have arisen by the
fragmentation of a normal chromosome (Kranz, 1971). B-chromosomes
suppress vegetative vigour and delay the onset of flowering (Jones,
1974), rare features observed, when compared with other members of Salvia
L.
The tetraploid member, - S. splendens Ker - Gawl. exhibit the highest TF%
value noted for the genus. Thus it can be c0nsidered.a~ primitive. Where
as, the diploid forms, S, coccinea Juss., - S. leucantha Cav. and &
plebeia R. Br. exhibit two SAT chromosomes each. Where as, the
tetraploid taxa, - S. splendens Ker-Gawl. exhibit four SAT-Chromosomes. In
plants, normally one satellited chromosome will be present in each basic
set of chromosomes (Bhaduri, 1944).
Adso~eles R. Br. In the two diploid members investigated in the genus
Anisomeles R.Br., A. indica (L.) Kuntze shows low values for all - -
karyomorphological characters. (vide Table 56; Fig.66) Eventhough the
lesser range of chromosome length and lesser variation coefficient value
denotes primitiveness, all the other characters including a low TF% (vide
Fig. 66, Table 56) shows the probale advanced status of A. indica (L.) - -
Kuntze than A- malabaica R. Br. The karyomorphological differences found
among these two taxa fully justify that speciation and evolution in this
genus has been principally effected by structural alteration of
chromosomes as observed in some other plants (Cleland, 1949).
Leucas R. Br. The different species of the genus w s R.Br. are
characterized by the homogeneous karyotypes as was evidenced by the
179
similar karyotype formula shown by all, except L, vestita Benth. Despite
the constancy in the chromosome number, the karyotypic differences among
the varieties were quite distinct (vide Table 56, Fig. 66).
In - L. vestita Benth., the high values obtained for all karyotypic details
except variation coefficient value (vide Fig. 66; Table 56) reveal it's
primitiveness, when compared with all the other cytotypes of Leucas R.Br.
Another remarkable feature shown by L, vestita Benth. is that they
contain only one pair of SAT chromosome when compared with two nucleolar
chromosomes found in & aspera Spreng., - L. cephalotes Spreng.,
linifolia Spreng. and - L. stricta Benth. (vide. Fig. 56). The secondary
constrictions are not of much importance when compared with other
karyotypic details, since they might be lost or gained during the course
of evolution (Lewis and John, 1963).
Leonotis R. Br. The higher karyomorphological values exhibited by L.
nepetifolia (L.) Ait.f. denotes it's primitive status. However the vast
difference of chromosome length and a high disparity index value (vide
Fig. 66; Table 56) denote an advanced heterogeneous nature. The
karyotypes that are heterogeneous both cytologically and genetically are
important in the evolution of species (Stebbins, 1958).
Teucrium L. In a more or less contradicting manner, the karyotype of 2
plectranthoides Gamble is characterized by a slight difference in
chromosome length as well as a high TF% value and hence seems to be
primitive. However, the low values for average chromosome length, total
chromatin length and chromatin length of basic complement (vide Fig. 66;
Table 56) point towards their evolved nature. Therefore the evolutionary
status of this taxa seems to be an enigma.
The karyomorphological diversity found in the different taxa of Lamiaceae
in South India can be best explained by assuming that the group is still
undergoing active speciation.
e) Meiotic Analysis
Meiotic studies are relevant, since they focus on details of pairing
behaviour of chromosomes, recombination frequencies and more importantly
their pattern of disjunction during Anaphase I and I1 which are not
deducible from mitotic studies (John and Lewis, 1965). Meiosis is an
integrated process consisting of a series of events which have been shown
to be under genetic control (Golubovskaya, 1979). Normal meiosis leads to
the formation of functional gametes, where as an irregular meiotic
division leads to abnormalities and there by to sterility (Del-Duca,
1976; Moraes - Fernandes, 1982).
In the present investigation, meiotic studies have been conducted on
twenty four members which bloom frequently in the experimental garden.
Out of these nineteen taxa exhibited normal meiosis and five taxa,
abnormal meiosis. Abnormalities in the normal course of meiosis,
especially failure of chromosome pairing has been reported to be
occurring spontaneously in various plant species (Gottschalk and Kaul,
1980).
Interestingly thirteen out of the nineteen taxa with normal meiosis are
polyploids. This might have been probably due to the absence of
multivalent associations (Swaminathan and Sulbha, 1959). The multivalent
formations which were found to be absent in Lamiaceae owe to the small
size of chromosomes. It is clear that well differentiated genomes are
involved in the origin of these polyploids (Koul and Gohil, 1991)
m b e r s with normal peiosis -
Out of the twentyfour taxa investigated, meiosis was perfectly normal in
six diploids and thirteen tetraploids. The diploids which exhibited
normal meiosis are Ocimum adscendens Willd., Orthosiphon glabratus
Benth., & glabratus var. parviflorus Benth., Salvia coccinea Juss., S,
leucantha Cav. and S. plebeia R. Br. -
The polyploids which exhibited normal meiosis included four varieties of
Ocimum basilicum L., four taxa of - 0. tenuiflorum L.f., two cultivars of
Orthosiphon grandiflorus Bold, & pallidus Royle, & thymiflorus Roth.
and Plectranthus wightii Benth. While considerable chromosomal
irregularities are expected in the metaphase I and anaphase I of this
plant, surprisingly it exhibits normal meiosis. This clearly indicates
the cytological stability attained by the polyploid to maintain it's
level. In all these species only bivalents are observed at diplotene and
metaphase I stages followed by normal disjunction of chromosomes to the
two poles.
This shows that irrespective of the degree of polyploidy, chromosomes
always exhibit bivalent formation. As a consequence polyploid meiosis is
characterized by bivalent formation. This can be used as an argument to
consider these plants as allopolyploids. However autopolyploids of long
standing tend to behave like allopolyploids (Giles and Randolph, 1951).
Another point of argument is that autopolyploids can undergo normal
meiosis as a result of the action of some genes which promote homologous
pairing only and not homeologous pairing (Sears and Okamoto, 1958).
mbers with abnormal meiosis
Abnormal meiotic behaviour is found in five out of the twenty four
members examined by smear experiments. The chances of survival of
polyploids are determined by their meiotic behaviour (Evans, 1988). The
tetraploids which exhibit abnormal meiotic status are Ocimum gratissimum
L., O, gratissimum L. var. suavis Hook. f., Plectranthus nummularius
Briq. and Salvia splendens Ker - Gawl. The hexaploid member - 0. americanum
L. also shows meiotic abnormalities. Meiosis in these polyploids are
characterized by the complete lack of multivalent associations even in
members with some what large chromosomes. This suggests that the size of
the chromosomes is perhaps no guide for multivalent formation, but it is
the genetic factors that controls chromosome pairing in autotetraploids
(Singh, 1991). This is evident from the discovery of genes which affect
different stages of meiosis (Rhoades and Dempsy, 1966)
The major meiotic abnormalities encountered in Ocimum americanum L., O,
gratissimum L. and 0- gratissimum L. var. suavis Hook. f. are the
occurrence of unpaired chromosomes at metaphase I and irregular
seggregation during anaphase I. This has been confirmed by earlier
observations made on the genus Ocimum L. (Sobti and Pushpangadan, 1982).
In Ocimum gratissimum L. and O, gratissimum L. var. suavis Hook. f. the
comparatively high number of univalents (vide Fig. 8j, 9j) reveal the
meiotic instability. A univalent is a chromosome that has not undergone
pairing at zygotene or may be the components of a bivalent separated at
diplotene owing to the absence of chiasma formation (Kihara, 1931;
Rosenberg, 1927). Thus the univalents at the end of first meiotic
division mechanically prevent complete separation of the two main groups
of chromosomes, leading to polyploidy (White, 1937).
In addition to this Ocimum gratissimum L. exhibits aberrant spores in the
form of pentads, hexads, octads etc. (vide Figs. 8n, 80, 8p). This may be
due to the unequal seggregation of chromosomes during telophase I1 or due
to the failure at cytokinesis in the last premeiotic division (Nath and
Nielson, 1963). Thus the occurrence of polyspory might be related to
chromosomal irregularities observed during meiosis (Longley, 1925).
The meiotic abnormalities found in Plectranthus nummularius Briq. (vide
Fig. 21j) might have forced the plant to resort to vegetative means of
reproduction, which was found to be the more prevalent method of
propagation. In many flowering plants exhibiting abnormal meiosis, escape
from the bottleneck of sterility is provided by the apomictic mode of
reproduction (Darlington, 1939).
The species Salvia splendens Ker - Gawl. exhibit partial or loose pairing
(vide Fig. 46j) Meiotic abnormalities were reported in many species of
Salvia L. by various authors (Delestaing, 1954; Linnert, 1955b; Haque and
Ghoshal, 1980a).
f) Cytological evolution ift Lamiaceae
From the cytological observations made in the present investigation it
can be concluded that speciation and evolution with in this family has
been possible as a result of increase in variability through changes in
the base numbers, as well as numerical and structural changes in
chromosome numbers.
The vnrious cytological phcnomenn like proto-auto pl.oidy, amphiploidy,
ascending and descending dysploidy might have resulted in the variability
of base numbers in the family (vide Fig. 67).
The wide range of chromosome numbers observed in many genera (vide
Fig.57) in the present investigation marks a significant role that an
aneuploidy and polyploidy have played in the evolution of various taxa of
the family at the generic and species level. It also appears that various
kinds of aberrations have played a vital role in the evolutionary
diversification of the family. The mitotic and meiotic irregularities
might have lead to structural and numerical variations in the chromosomes
of a species. Induviduals with the same chromosome number but with
differences in karyomorphological details reflect the ongoing
evolutionary processes at micro level.
It seems probable that in South Indian Lamiaceae, Robertsonian changes or
mutations might have also played an important role in the evolution of
karyotype. Drastically mutated induviduals are usually unstable and unfit
to survive in nature because they express various degrees of weakness and
chromosomal aberrations leading to genetic sterility. However some
induviduals carrying the changed chromosomal constitutions are well
within their range of tolerance. This was confirmed by the occurrence of
normal meiosis in some polyploids investigated.
More over, most of the taxa belonging to this family have efficient means
of vegetative propagation. This ensures the survival of these genetically
altered types which otherwise would have faced extinction on account of
sexual sterility imposed as a result of these changes. Accumulation of
such small changes can sometimes lead to a taxonomic divergence in a
species during the process of evolution. Thus meiotic accidents may prove
to be more useful than mitotic aberrations from the evolutionary point of
view, since the meiotic abnormalities are hereditary and likely to
multiply and establish in a population.
Polyploidy is said to have an ancient origin if a taxon includes only
polyploids or very few diploids, and relatively of the recent origin if
diploids are more frequent than polyploids or both are in almost equal
proportion (Stebbins, 1950). On this context among the members
investigated, the various genera, viz., Ocimum L., Orthosiphon Benth.,
Plectranthus L' Herit., Coleus Lour., Hyptis Jacq. and Mentha L.,
polyploidy is assumed to have an ancient origin. However polyploidy seems
to be of recent origin in Pogostemon Desf., Salvia L., Anisomeles R.Br.
and Leucas R.Br. An effective conclusion cannot be reached in the genera
like Acrocephalus Benth., Eusteralis Rafin., Leonotis R.Br. and Teucrium
L., since only one taxa has been investigated in each of them.
Thus the family Lamiaceae seems to be in a fairly active state of
evolution because of the quite common occurrence of polyploidy and
aneuploidy. Mutation of genes, structural changes in chromosomes, non-
disjunction, chromosomal rearrangement and several other abnormalities
are those mechanisms leading to the development of polyploidy and thus to
the differentiation of new taxa in Lamiaceae (Fernandes and Leitao,
1984).
There are still enormous gaps in our knowledge as regards the cytological
evolution of Lamiaceae, and much still remains to be done before a major
cytotaxonomic review may be attempted.
Fifty four taxa representing a total of fifteen genera of Lamiaceae found in
South India are examined cytologically. The chromosome spectrum in South
Indian Lamiaceae ranges from 2n=16+0-1 B to 2n596, with majority of the
species concentrated in the number 2n = 32, followed by 2n = 48, 2n = 28 and
2n = 22. In spite of the wide range of chromosome numbers, there exists a
relationship between the different genera and species as evidenced by the
frequent occurrence of numerical variations. This wide range of chromosome
numbers may be due to the difference in number of chromosomal biotypes
belonging to the different groups. In the present investigation, the genera
like, Ocimum L., and Coleus Lour. exhibit inter, intra and infra specific
variations among chromome numbers. Both inter and intraspecific variations
have been found in Anisochilus Wall., Hyptis, Jacq., Pogostemon, Desf., Mentha
L. and Salvia L. Where as, Orthosiphon Benth., exhibit both inter and infra
specific variations. Inter specific variations alone has been a characteristic
feature shown by Plectranthus L'Herit., Anisomeles R.Br. and Leucas R.Br. The
presence of such widely different series of chromosome numbers in the species
of even the same genus and in genera placed under different tribes and
subtribes, indicate that the different chromosome numbers may be derived one
from the other.
The basic chromosome numbers are found to be widely varied in South Indian
Lamiaceae. This great variability in the number of chromosomes at the basic
level could possibly be the result of aneuploidy at generic level. Out of the
fifteen genera investigated, three (Ocimum L., Orthosiphon Benth. and Coleus
Lour.) have been found to be polybasic, five (Plectranthus L' Herit.,
Anisochilus Wall., Hyptis Jacq., Salvia L. and Anisomeles R.Br.) to be dibasic
and three (Pogostemon Desf. Menthn L. and Leucas R.Br.) to be monobasic. As
the following genera, viz. Acrocephalus Benth., Eusteralis Rafin., Leonotis
R.Br. and Teucrium L. are represented by only one of their species in the
present study, it would not be appropriate to classify them as monobasic. Both
primary as well as secondary base numbers are found to be involved in the
evolution of the fiftyfour taxa investigated. The primary base numbers (Xl)
range from 6 to 9 and secondary base numbers (X2) from 10 to 13 and 15 to 17.
The basic chromosome number X2 a 12 is prevalent in majority of the members
(24.07%). This is followed by the base numbers X2 = 11, X1 = 7, X1 = 8 and X2
= 16 with percentages 16.67, 14.81, 12.96 and 11.11 respectively. Thus it is
likely that protopolyploidy plays an active role in the evolution.
The family Lamiaceae is characterized by a relatively high frequency of
polyploidy. Studies conducted on fifty four taxa reveals the dominance of
polyploids (62.96%) over diploids in South Indian Lamiaceae. Out of these,
57.41% were found to be tetraploids followed by hexaploids (3.7%) and
octaploids (1.85%). The present study also reveals that the herbaceous
elements predominate the shrubby elements of the family. Correlating
polyploidy with life form and growth habit, it is clear that the highest
frequency of polyploids occur in perennial herbs (79.41%) lowest in annual
herbs and undershrubs (2.94% each) and intermediate in perennial undershrubs
(14.71%). The various genera which show predominance of polyploidy include
Ocimum L., Orthosiphon Benth., Coleus Lour., Hyptis Jacq., Pogostemon Desf.,
Mentha L., Salvia L., Leonotis R. Br. and Teucrium L. The role of both
polyploidy and aneuploidy in the mechanism of speciation is obvious in South
Indian Lamiaceae. *
The general feature noted in the family Lamiaceae is the wide range of
chromosomes with graded symmetrical karyotypes. However the chromosome
complements in South Indian Lamiaceae members differ in minute karyotypic
details. With regard to gross morphology chromosomes are nearly submetacentric
in nature. The chromosomes were found to range from 4 . 2 p to 1.0 ,u in length.
The chromosomes with secondary constriction range from two to six in number.
The average chromosome length varies from 2 . 7 4 ~ to 1.12 p. The total
chromatin length shows a very wide variation with 2 4 . 4 ~ being the minimum and
154.8 p being the maximum value. The disparity index values were found to
range between 22.22 and 5 0 . The coefficient of variation ranges from 12.53 to
30 .00 and total centromeric index (TF%) from 35 .83 to 45 .07 . Thus the various
micromorphological details of the karyotype like, differences in absolute
chromosome size, differences in the position of centromere, differences in
total chromatin length, differences in karyotype formula and differences in
the number as well as position of satellites vary from cytotype to cytotype.
These variations found in the karyotypic parameters suggest that South Indian
Lamiaceae members are characterized by symmetrical to slightly asymmetrical
karyotypes. The karyomorphological diversity found in the family shows that
the group is still undergoing active speciation.
Meiotic studies have been conducted on twenty four taxa which frequently
blossom in the experimental garden. Out of these, nineteen taxa exhibited
normal meiosis and five taxa, abnormal meiosis. Interestingly thirteen out of
the nineteen members with normal meiosis are polyploids. Multivalent
associations are found to be invariably absent in the taxa examined. The
absence of multivalent formations owe to the small size of chromosomes. Thus
it is clear that well differentiated genomes are involved in the origin of
these polyploids. It has also been found that irrespective of the degree of
polyploidy, chromosomes always exhibit bivalent formation.Polyploid meiosis
characterized by bivalent formation point towards allopolyploidy. However
autopolyploids on long standing behaves like allopolyploids. The major
aberrations encountered in members which exhibit abnormal meiosis are the
occurrence of unpaired chromosomes at metaphase I, irregular segregation
during anaphase I and the disposition of partial or loose pairing. These
abnormalities reveal the meiotic instability leading to polyploidy in many
taxa of the family.
The presence of a wide range of chromosome numbers, numerical variations and
structural changes of chromosomes found in many genera mark the significant
role that both aneuploidy and polyploidy have played in the evolution of
various taxa of the family at the generic and species level. The variability
in base numbers might have been resulted through proto autoploidy,
amphiploidy, ascending and descending dysploidy. Both mitotic and meiotic
aberrations have played a major role in the evolutionary diversification of
the family. Induviduals with same chromosome number but with differences in
karyomorphological details reflect the ongoing evolutionary processes at
microlevel. It has also been found that in South Indian Lamiaceae,
Robertsonian changes or mutations might have also played a major role in the
evolution of karyotype.